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ISSN 0968-0454 

Bulletin of 
The Natural History Museum 


Ex I O 

ntomology Series 

23 AUG 2uii2 

®RN£«AL LtSRAflv 

VOLUME 71 NUMBER 1 27 JUNE 2002 

The Bulletin of The Natural History Museum (formerly: Bulletin of the British 
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The Entomology Series is produced under the editorship of the 
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World list abbreviation: Bull. nat. Hist. Mus. Lond. (Ent.) 

ISSN 0968-0454 

The Natural History Museum Entomology Series 

Cromwell Road Vol. 71, No. 1, pp. 1-133 

London SW7 5BD Issued 27 June 2002 

Typeset by Ann Buchan (Typesetters), Middlesex 

Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset 


Bull. nat. Mas. Land. (Ent.)71(l): 1-18 

Issued 27 June 2002 

Loboponera gen. n. and a review of 
the Afrotropical Plectroctena genus 
group (Hymenoptera: Formicidae) 


Department of Entomology, The Natural History Museum, Cromwell Road. London 
SW7 5BD, UK 



The tribe Ponerini 

The Plectroctena genus group 

Loboponera gen. n 

List of species 

Key to species 

Synopsis of species 

Psalidomyrmex Andx6 

List of species 

Key to species 




Synopsis of species 9 

Plectroctena F. Smith 10 

List of species 10 

Key to species 1 1 

Synopsis of species 12 

References 15 

SYNOPSIS. The Afrotropical ponerine ant genus Loboponera is newly described, ll contains 7 
new species and one species previously regarded as a member of Pachycondyla F. Smith (Z.. nasica 
(Santschi) comb. n.l. The relationships of Loboponera within the Plectroctena genus group and 
tribe Ponerini are discussed and apomorphy-based diagnoses are provided for all taxa. An 
identification key and descriptions are provided for the workers of Loboponera. Re\ ised keys and 
taxonomic synopses are given for Psalidomyrmex(6 species ) and Plectroctena ( 1 5 species ); within 
the latter PI. gabonensis is a new junior synonym of PI. minor. 


The name Loboponera first appeared in print in Bolton's 
(1994: 1 56) key to the Afrotropical Ponerinae. where it 
was taxonomically unavailable. The genus was known 
to be undescribed several years earlier and was in- 
cluded so that a full picture of the regional fauna could 
be provided, using the name assigned to it by William 
L. Brown Jr. At that time Brown was engaged in a 
revision of the world Ponerini genera and the appear- 
ance of a formal description for Loboponera was 
expected soon afterwards. Sadly, Professor Brown died 
before he had completed his studies and so his work on 

Loboponera, and the rest of the Ponerini, went into 

More recently Steven O. Shattuck ( ANIC. Canberra) 
has retrieved Brown's Ponerini manuscripts and is 
currently working on them in an attempt to bring the 
study of the genera to a conclusion. He passed on 
Brown's Loboponera notes to me, so that I could 
conclude work on this genus to species rank. Stefan 
Cover (MCZ, Harvard) kindly provided me with the 
figures for the genus that Brown had prepared and they 
are published here. 

While studying material of the Loboponera species 
it became apparent that they formed a monophyletic 
group with two other Afrotropical Ponerini genera, 

© The Natural History Museum, 2002 


Psalidomyrmex and Plectroctena, both of which had 
been revised previously (Bolton, 1974, 1975). This 
presented a reasonable opportunity to update our know- 
ledge of these two genera and at the same time introduce 
the new genus, so that a group diagnosis and new 
findings for all of them could be included in a single 
publication. Diagnoses of all three genera are presented, 
as are comparative notes and keys to species. Formal 
diagnoses of the Loboponera species are given but the 
species of the other two genera are more summarily 
treated as revised descriptions already exist. Thus for 
the species of Psalidomyrmex and Plectroctena only 
taxonomic synopses are presented, together with addi- 
tional distribution records. 

It should be pointed out that a subtribal name, 
Plectroctenina, exists for this group. It was originally 
spelled Plectroctenini Emery, 191 1 : 92, but the suffix 
is modified here in accordance with the Fourth Edition 
of the International Code of Zoological Nomenclature 
(1999). At present this subtribal name is treated as a 
junior synonym of Ponerini (Bolton, 1994, 1995). As 
well as Plectroctena and Psalidomyrmex, Emery also 
included in his subtribe Myopias Roger and 
Trapeziopelta Mayr, the latter of which is now a junior 
synonym of the former (Willey & Brown, 1983: 249). 
Myopias, distributed through the Oriental, Malesian 
and Austral regions, is not part of the Plectroctena 
group. Its affinities appear to lie with Pachycondyla 
and its immediate allies. 

The three genera of the Plectroctena group are 
strictly Afrotropical, with a majority of species in the 
forest zones of West and Central Africa. All nest in 
rotten wood or in the soil and foraging is carried out 
there and in the leaf litter layer. Some species may also 
use abandoned termitaries as nest sites (Dejean, Durand 
& Bolton, 1996). Diet where known appears entirely 
carnivorous and the species are specialised predators. 
Plectroctena species prey mostly, or perhaps in some 
species entirely, on millipedes or their eggs (Levieux 
(1972), Bolton (1974), Bolton, Gotwald & Leroux 
( 1 979)). PI. minor has been recorded as preying on 
termites as well as millipedes (Dejean & Suzzoni, 
1 99 1 ). Arnold (1915) mentions millipedes, beetles and 
termites forming the diet of PL mandibular is, though 
Hamish Robertson (SAM, personal communication) 
informs me that the record for beetles as prey has never 
been confirmed. Psalidomyrmex procerus captures 
earthworms (Levieux (1982), Dejean, Schatz, Orivel 
& Beugnon (1999)); the prey of Loboponera remains 
unknown. Little detail is known of the biology 
and ecology of individual species apart from 
comments in the publications just mentioned and some 
scattered remarks in the literature of original descrip- 
tions, but Villet (1991) has reported on colony 
foundation by, and the origin of, ergatoid queens in PI. 
mandibular is. 

Standard measurements and indices, and abbrevia- 

tions for the names of museums, are as noted in the 
earlier publications (Bolton, 1974. 1975). 

The tribe Ponerini 

The subfamily Ponerinae, as it is presently understood, 
is probably paraphyletic. Bolton (1990a, 1990b) and 
Baroni Urbani, Bolton & Ward (1992) proposed 
tergosternal fusion of the fourth abdominal segment as 
a synapomorphy of the subfamily, but the validity of 
the character was compromised by Ward (1994). In 
consequence the tribal composition, and indeed the 
monophyly, of the Ponerinae is still very much in 
doubt (Keller, 2000). 

Of the Ponerinae tribes that are currently recognised, 
with varying degrees of confidence, the Ponerini is 
easily the largest (Bolton, 1994). In recent years tribe 
Amblyoponini has been redefined by Ward (1994) and 
the phylogeny of the Ectatommini and its relatives has 
been investigated by Lattke (1994) and Keller (2000). 
At present the tribe Ponerini is diagnosed by the 
following synapomorphic characters. 

1 . Median portion of clypeus is narrowed posteriorly 
and is narrowly inserted between the frontal lobes 
as a slender triangle or linear strip (worker and 

2. Inner borders of frontal lobes are very closely 
approximated or confluent for most or all of their 
length, often separated merely by a sulcus (worker 
and queen). 

3. Outer borders of frontal lobes form simple short 
semicircles or blunt triangles and have a distinctly 
pinched-in appearance posteriorly (worker and 

4. Helcium projects from low down on the anterior 
face of the first gastral (= third abdominal) segment 
and the anterior face above the helcium is vertical 
and usually high (worker and queen). 

5. Mandibles are reduced, short to lobate, edentate 
except for apical tooth at most (male). 

The Plectroctena genus group 

Within the tribe Ponerini a small group of Afrotropical 
genera, Plectroctena + Psalidomyrmex + Loboponera, 
share the following autapomorphic development in 
workers and queens. 

Anteroventral articulators surface of petiole long 
and very broad, the surface with a narrow median V- 
shaped longitudinal groove or central small pore-like 
depression (see comments for discussion). 

Other characters that in combination are diagnostic 
of female castes in the group include the following. 
Some are plesiomorphic but some (marked with an 
asterisk) are probable apomorphies of which analogues 
have apparently developed convergently elsewhere in 
tribe Ponerini. 


*Frontal lobes hypertrophied. 

Antenna with 12 segments. 

Promesonotal suture strongly developed. 

Orifice of propodeal spiracle round or very broadly 

oval, nearly round. 
Metapleural gland orifice lateral. 
Metanotal groove usually absent, rarely vestigial. 
*Mesotibia and metatibia each with only a single spur 

(also in males). 
Pretarsal claws simple, small. 
Petiole nodiform in profile. 
Constriction between first and second gastral segments 

deep and broad. 
*Stridulitrum absent from pretergite of second gastral 

(= 4th abdominal) segment. 


Within Ponerini but outside this group the anteriormost 
part of the ventral surface of the petiole, where it 
articulates with the alitrunk, is quite uniform. At the 
junction, immediately behind the alitrunk margin and 
intersegmental membrane, the petiole has an arched or 
broadly horseshoe-shaped strip of cuticle which, when 
the petiole is flexed down, slides into the alitrunk: this is 
the articulating surface proper. Immediately behind this 
is a transverse impression or depressed area which is 
followed by a flat or slightly convex zone upon which 
there are usually a number of proprioceptor hairs. 

The members of the Plectroctena genus group have 
modified this basic arrangement by extending the length 
of the articulating surface of the petiole posteriorly on 
each side, and expanding its width towards the ventral 
midline. This has the effect of constricting the subse- 
quent areas into a narrow median impression or groove. 
In most members of the group this groove is a narrow 
inverted V-shape but in some (Plectroctena lygaria, 
Psalidomyrmexfoveolatus) the development has gone 
so far that only a tiny median pore-like depression 

Formal diagnoses of the three genera included in the 
group are given below, but they can be distinguished 
quickly in the worker and queen castes by the follow- 
ing features. 

Plectroctena has linear mandibles and a large semi- 
circular excavation in the clypeal margin at the 
mandibular articulation. 

Psalidomyrmex has the labrum projecting as a promi- 
nent lobe in front of the anterior clypeal margin and the 
mandible has a long attenuated apical tooth. 

Loboponera has the median portion of the clypeus 
produced into a lobe that overhangs the mandibles and 
has the second gastral tergite strongly vaulted. 

Dolioponera Brown (1974) may also belong to this 
genus group but as the taxon is known only from a 
single worker, the holotype, there is no chance of 
dissection to assess the presence of the apomorphy. 
Nevertheless, it has most of the features noted above 

and the mandible, with its rounded basal angle, is 
reminiscent of Psalidomyrmex. There are however 
some striking differences. Dolioponera lacks the 
characteristic labral lobe of Psalidomyrmex, the me- 
dian portion of its clypeus projects as a short truncated 
lobe, the propodeum lacks a median longitudinal groove 
and the second gastral segment is elongate and cylin- 
drical. Finally, the three genera of the group all have an 
anteroventral process on the first gastral stemite that 
projects somewhat forward and downward beneath the 
helcium; this is absent in Dolioponera. 

LOBOPONERA Bolton & Brown gen. n. 

(Figs. 1-11) 

[Loboponera Bolton, 1994: 156 (in key), unavailable 

Type-species: Loboponera vigilans, by present desig- 

Worker. Ponerine ants with the tribe and group 
characters noted above and also with the following. 

1 . Median portion of clypeus extended anteriorly as a 
lobe that projects out over the mandibular basal 
margins: anterolaterally the clypeus with a small 
projecting lobe that overhangs the outer base of the 

2. Labrum not visible in full-face view with mandi- 
bles closed. 

3. Mandible subtriangular to triangular; apical tooth 
the largest but not attenuated; basal angle narrowly 
rounded to angulate. 

4. Palp formula 2,2 (dissections of obeliscata, 

5. Posteroventral curve of head in profile with a 
projecting curved flange that is formed from the 
hypertrophied lateral portion of the occipital carina. 

6. Propodeal dorsum without a median longitudinal 
groove or impression. 

7. Mesofemur and metafemur with a longitudinal 
(glandular?) groove present mid-dorsally on basal 

8. Tergite of second gastral (= 4th abdominal) segment 
vaulted, strongly arched and down-curved 
posteriorly; second gastral sternite correspondingly 

QUEEN. The few known (basalis. nasica, vigilans) 
are slightly larger than the worker, dealate, with the 
above characters and the usual modifications associ- 
ated with this caste. 

Male. Unknown. 


Characters 1, 4, 5 and 8 are autapomorphic; 2, 3 and 6 
are plesiomorphic. Character 7 is equivocal; it may be 

autapomorphic but synapomorphy with Plectroctena, 
which also has femoral grooves developed, is a possi- 
bility. The latter case would leave Psalidomyrmex, 
which has no trace of femoral grooves, either 
plesiomorphic for this character or apomorphic by 
secondary loss. 

Species of Loboponera superficially resemble the 
Afrotropical members of Pachycondyla that are stockily 
built, more heavily sculptured and have a nodiform 
petiole (i.e. former members of Bothroponera). 
Indeed, the only previously described species, nasica, 
was originally included there. However, those 
Pachycondyla species (in the broad sense of Bolton, 
1994, 1995) that superficially resemble Loboponera, 
beside lacking the petiole articulation autapomorphic 
in the Plectroctena group, also differ as they have the 
following features. 

Mesotibia and metatibia each with two spurs, one large 
and pectinate, the other smaller and simple. 

Mesofemur and metafemur without a mid-dorsal lon- 
gitudinal groove. 

Posteroventral curve of head without a projecting 

Stridulitrum present on pretergite of second gastral 

Tergite of second gastral segment curved but not 

Orifice of propodeal spiracle usually slit-shaped. 

Palp formula usually greater than 2,2. 

Loboponera also shows some striking parallelisms 
with the ectatommine genus Gnamptogenys. Their 
habitus similarities may indicate that Loboponera 
shares some ecological or behavioural features with 
the latter genus. This is an interesting speculation as 
Gnamptogenys, with many species in the Neotropical, 
Oriental and Malesian regions (Brown, 1958; Lattke, 
1995 and in preparation), is absent from the 
Afrotropical; Loboponera on the other hand is re- 
stricted to the Afrotropical region. All collections of 
this genus have been recovered from leaf litter, rotten 
wood or abandoned termitaries. 

List of species 

obeliscata group 

obeliscata Bolton & Brown sp. n. 

politula Bolton & Brown sp. n. 
trica group 

trica Bolton & Brown sp. n. 
vigilans group 

basalis Bolton & Brown sp. n. 

edentula Bolton & Brown sp. n. 

nasica (Santschi, 1920) comb. n. 

subatra Bolton & Brown sp. n. 

vigilans Bolton & Brown sp. n. 


Key to species (workers) 

1 Dorsum of alitrunk and first gastral tergite without stand- 
ing hairs of any form (Figs. 4—6). Second gastral tergite 
punctate or with fine dense sculpture but without longitu- 
dinal costate sculpture. With head in full-face view the 
sides without projecting hairs (Figs. 2-3) 2 

- Dorsum of alitrunk and first gastral tergite with numerous 
to abundant stiff, usually short, standing hairs (Figs. 7-9). 
Second gastral tergite with coarse longitudinal costate 
sculpture. With head in full-face view the sides with at 
least 1-2 sho-rt projecting hairs (Fig. 1 ) 4 

2 Leading edge of scape without projecting hairs; dorsum 
of scape finely and densely punctulate. Dorsal surfaces of 
head and body, including the frontal lobes, entirely 
blanketed with very fine dense sculpture, opaque. 
Propodeal lamella restricted to lower half of declivity. 
(Ivory Coast) trica 

- Leading edge of scape with a row of suberect projecting 
hairs; dorsum of scape longitudinally costulate-rugulose. 
Dorsal surfaces of head and body not entirely finely 
densely sculptured, not opaque. Propodeal lamella extends 
the whole depth of the declivity 3 

3 With petiole in profile the dorsum rising medially to a 
bluntly subpyramidal point (Fig. 4). Scape longer, SI > 
95; when laid back the apex of the scape just reaches the 
occipital margin (Fig. 3). Larger species, HW > 0.90. 
(Ghana, Gabon) obeliscata 

- With petiole in profile the dorsum shallowly convex to 
flat, not rising to a median subpyramidal point (Fig. 5). 
Scape shorter, SI < 90; when laid back the apex of the 
scape distinctly fails to reach the occipital margin (Fig. 
2). Smaller species, HW < 0.70. (Ivory Coast, Ghana. 
Nigeria) politula 

4 First gastral sternite with a mid-ventral longitudinal 
cuticular crest or carina that may extend entire length of 
sternite or be restricted to posterior half; carina usually 
partially to entirely translucent and clearly visible in 
profile 5 

First gastral sternite without a mid-ventral longitudinal 
cuticular crest or carina that is partially to entirely trans- 
lucent and clearly visible in profile 7 

5 Base of gaster deeply concave when viewed from above 
and slightly to the front; on each side the tergite at its 
lowest point, at about the level of the helcium. projects 
forward and curves medially, forming a strongly defined 
incurved lobe (Figs. 9-10). (Ivory Coast, Ghana, Camer- 
oun) basalis 

- Base of gaster transverse to shallowly concave when 
viewed from above and slightly to the front; on each side 
the tergite at its lowest point, at about the level of the 
helcium. forms a simple angle and does not project 
forward and curve medially as an incurved lobe 6 

6 In dorsal view first gastral tergite coarsely foveolate, not 
costulate; spaces between foveolae raised and flat, entirely 
covered with fine dense shagreenate sculpture, dull and 
opaque. Longitudinal costae of second gastral tergite 


densely shagreenate and opaque. Eye large, with > 10 
ommatidia in its maximum diameter and that diameter 
greater than maximum width of second funicular seg- 
ment (Fig. 7). (Ivory Coast. Nigeria, Cameroun, Gabon, 
Zaire) vigilans 

In dorsal view first gastral tergite predominantly longitu- 
dinally costulate or rugulose, with scattered foveolate 
punctures between the costulae/rugulae; entire surface 
glossy, without dense opaque shagreenate sculpture. 
Longitudinal costulae of second gastral tergite glossy, not 
densely shagreenate and opaque. Eye small, with < 10 
ommatidia in its maximum diameter and that diameter 
less than maximum width of second funicular segment 
(Fig. 8). (Ivory Coast, Ghana, Cameroun, Gabon) 

1 Propodeal declivity with a small tubercle above a larger 
more basally situated tooth. Side of head behind and 
below eye sculptured with large, broad-rimmed coarse 
foveolate punctures, not costulate. With head in profile 
the lower occipital curve with a large lobate cuticular 
flange that is much broader than the maximum diameter 
of the eye. (Cameroun) subatra 

- Propodeal declivity unarmed and smoothly rounded above 
a basally situated tooth. Side of head behind and below 
eye sculptured with fine longitudinal costulae and 
scattered small foveolate punctures. With head in profile 
the lower occipital curve with a small shallow cuticular 
crest that is narrower than the maximum diameter of the 
eye. (Rwanda) edentula 

Synopsis of species 

obeliscata group 

(Figs. 2-5) 

Mandible with 1-2 teeth between apical tooth and 
basal angle. 

Median portion of clypeus, anterior to frontal lobes, 
narrow but not strongly bilaterally compressed; with- 
out a median longitudinal carina. 

Propodeal declivity with a simple lamella on each side 
that extends its entire height; without a projecting 
lobe or tooth close to the base of the declivity. 

Sculpture between punctures on dorsal alitrunk and 
gaster mostly to entirely effaced; surfaces between 
punctures mostly to entirely smooth and shining. 

Dorsal surfaces of head behind frontal lobes, alitrunk, 
petiole node and first gastral tergite without stand- 
ing hairs; second gastral tergite with standing hairs 
restricted to apical quarter. 

Scape with projecting hairs on the leading edge. 

Loboponera obeliscata sp. n. 

(Figs. 3, 4) 

HOLOTYPE WORKER. TL5.6,HL 1.13, HW 1.00, CI 
88, SL 0.97, SI 97, AL 1 .64. Characters of obeliscata 
group and the following. Eye minute and difficult to 

see, scarcely larger than one of the adjacent foveolate 
punctures; maximum diameter 0.04. Scape when laid 
back in full-face view reaches occipital margin. Small 
lobe at anterolateral angle of clypeus distinctly promi- 
nent. Propodeal lamella broad at sides, narrowing near 
dorsal angle and thinly continued onto dorsum but 
narrowed medially or with a mid-dorsal gap. Propodeal 
declivity finely transversely striate. Petiole in profile 
with anterior and posterior margins more or less paral- 
lel and with the dorsal surface extended medially into 
a high subpyramidal blunt point; in posterior view the 
sides slope steeply upward to a pointed apex. First 
gastral tergite in profile with its anterior face distinctly 
inclined posteriorly from top to bottom, in dorsal view 
overhanging the helcium. 

Paratype worker. TL 5.7, HL 1 . 14, HW 1 .00, CI 
88, SL 1.00, SI 100.AL 1.72. 

Holotype worker. Ghana: Tafo, 9.ix. 1 966, ant ecol- 
ogy sample 251 (D. Leston) (BMNH). 

Paratype. 1 worker with same data as holotype 

Non-paratypic material examined. Gabon: La 
Makande, Foret des Abeilles (A Dejeari). 

Closest related to politula but easily distinguished 
by the unique shape of the petiole node. 

Loboponera politula sp. n. 

(Figs. 2, 5) 

Holotype worker. TL 3.7, HL 0.73. HW 0.64, CI 
88, SL 0.52, SI 81, AL 1.09. Characters of obeliscata 
group and the following. Eye minute, maximum diam- 
eter 0.03. Scape when laid back in full-face view 
distinctly fails to reach occipital margin. Propodeal 
lamella narrow at sides, petering out at dorsal angle 
and not continued onto dorsum; the latter separated 
from the declivity by an angle. Petiole in profile with 
anterior margin rising to a blunt anterodorsal angle, 
behind which the dorsum slopes posteriorly and is flat 
or very feebly convex. In posterior view the petiole 
node dorsum is broadly evenly rounded. First gastral 
tergite in profile or in dorsal view with its anterior face 
overhanging the helcium. 

Paratype workers. TL 3.4-3.7, HL 0.70-0.73, HW 
0.60-0.64, CI 85-90, SL 0.48-0.52, SI 79-82, AL 
1.01-1.09 (3 measured). 

Holotype worker, Nigeria: Ibadan, IITA no. 62, 
16.vii.1974 (B.R. Critchley) (BMNH). 

Paratypes. 1 worker, Ivory Coast: Abidjan, Banco 
Nat. Pk. 18.iii.1977, primary forest litter (/. Lobl)\ 1 
worker, Ghana: Mampong, 10.viii.1970 (P. Room); 1 
worker, Nigeria: near Ibadan, 26.vii.1981, secondary 
forest, no. 10 (A. Russell-Smith) (BMNH, MCZ). 

Non-paratypic material examined. Ghana: Bunso, 
nr Tafo (R. Belshaw). 

This species is smaller than the above and, beside 


lacking the unique subpyramidal shape of the petiole 
node that characterises obeliscata, has much shorter 

trica group 

(Fig. 6) 

Mandible with 3 teeth between apical tooth and basal 

Median portion of clypeus, anterior to frontal lobes, 
strongly bilaterally compressed, narrow and with a 
nearly vertical longitudinal carina. 

Propodeal declivity with a rounded lobe near base, 
without a lamella above the lobe. 

Sculpture between punctures everywhere on dorsal 
head and body fine and very dense. 

Dorsal surfaces of head behind frontal lobes, alitrunk, 
petiole node and first gastral tergite without stand- 
ing hairs; second gastral tergite without standing 

Scape without projecting hairs on the leading edge. 

Loboponera trica sp. n. 

(Fig. 6) 

Holotype worker. TL 3.0, HL 0.59, HW 0.55. CI 
93, SL 0.43, SI 78, AL 0.90. Characters of trica group 
and the following. Maximum diameter of eye 0.02. 
Scape when laid back not reaching occipital margin. 
Dorsum of head opaque, extremely densely finely 
striolate-punctulate and with larger shallow foveolae 
that are overlaid and partially masked by the finer 
components of the sculpture. Alitrunk and gaster simi- 
larly sculptured, in places with the foveolae more 

Paratype worker. TL 3.0, HL 0.59, HW 0.55, CI 
93, SL 0.47, SI 85, AL 0.90. 

Holotype worker, Ivory Coast: Abidjan, Banco Nat. 
Pk, 14.iii.1977, dead wood sample (/. Lobl) (BMNH). 

Paratype. 1 worker with same data as holotype 

L. trica is a very distinctive small species that is 
easily recognised by its lack of standing hairs coupled 
with the presence everywhere of uniform fine dense 

vigilans group 

(Figs. 1,7-11) 

Mandible with 3-5 teeth between apical tooth and 

basal angle. 
Median portion of clypeus, anterior to frontal lobes, 

usually with a low median longitudinal carina. 
Propodeal declivity with a lobe or triangular tooth near 

base; margin above this either unarmed, or with a 

narrow lamella, or with a small tooth, or with a 

lamella and a tooth. 

Sculpture coarse and dense on dorsal alitrunk and 
gaster, basically of coarse foveolae with rugulose or 
costate-striate interspaces. 

Dorsal surfaces of head behind frontal lobes, alitrunk, 
petiole node and first gastral tergite with numerous 
short suberect to erect hairs; second gastral tergite 
with standing hairs everywhere. 

Scape with projecting hairs on the leading edge. 

Loboponera basalis sp. n. 

(Figs. 1,9, 10) 

Holotype worker. TL 6.6, HL 1 .20, HW 1 .06, CI 
88, SL 1.02, SI 96, AL 1.87. Characters of vigilans 
group and the following. Maximum diameter of eye 
0.07. First gastral sternite with a mid- ventral longitudi- 
nal translucent carina that extends the length of the 
segment. First gastral tergite in profile with its 
anteroventral angle extended forward as a broadly 
triangular tooth, just above the helcium. In ventral 
view the extended anteroventral angles of the first 
tergite are seen to be drawn out as incurved triangular 
broad teeth or short stout horns on each side of the 
helcium. In dorsal view the anterior face of the first 
gastral tergite forms a broad concave arc that is extended 
forward and inward on each side by the broadly 
triangular curved angles. Propodeal declivity above 
the basal tooth with a narrow uneven lamella that 
terminates dorsally in a small tooth or tubercle. 
Sculpture everywhere on the glossy dorsum longitudi- 
nally costate-rugulose between broad foveolate 
punctures that are mostly sharply defined. On the 
second gastral tergite the longitudinal costate-rugose 
sculpture predominates. 

Paratype workers. TL5.9-6.6,HL1.11-1.16,HW 
1.05-1.11, CI 95-96, SL 1.00-1.08, SI 94-97, AL 
1.82-2.02 (3 measured). 

Holotype worker. Ivory Coast: Lamto, Toumodi, 
5.iii.l968, rotten wood (J. Levieux) (MCZ). 

Paratypes. 2 workers with same data as holotype; 1 
worker. Ivory Coast: Nzi Noua, on Ndouci-Toumodi 
Highway, about 21 km. N. of Ndouci, in rotten wood 
(J. Levieux); 1 worker, Ghana: Tafo, rotten wood (D. 
Leston); 2 workers, Tafo, 25. v. 1970, on ground at base 
of tree (B. Bolton) (MCZ, BMNH, MNHN). 

Non-paratypic material examined. Ivory Coast: 
Iringou (F. Krell). Cameroun: Kala (A. Dejean). 

The unique structure of the base of the first gastral 
tergite immediately isolates this very conspicuous spe- 

Loboponera edentula sp. n. 

Holotype worker. TL 6. 1 , HL 1 . 1 1 , HW 1 .03, CI 
93, SL 0.98, SI 95, AL 1.78. Characters of vigilans 
group and the following. Maximum diameter of eye 


0.07. First gastral sternite without a tooth on each side 
of the anteroventral median process and without a 
translucent median longitudinal carina. Propodeal 
declivity with a small sharp tooth near base but other- 
wise unarmed and rounding evenly into the dorsum. 
Sculpture on glossy head and alitrunk is of relatively 
fine longitudinal costulae and scattered foveolate punc- 
tures. On the first gastral tergite the costulae are mostly 
supressed, much less obvious than on the alitrunk, and 
the spaces between the foveolae are very glossy and 
partially to mostly smooth. 

Paratype worker. TL 6. 1 , HL 1 . 1 1 , HW 1 .04, CI 
94, SL 0.94, SI 90, AL 1.79. 

Holotype worker, Rwanda: Kayove, 23. iv. 1973, 
2100 m (P. Werner) (MCZ). 

Paratype. 1 worker, Rwanda: Rangiro, ix. 1976, litter 
(P. Werner) (BMNH). 

The form of the propodeal declivity, coupled with 
the lack of a longitudinal carina on the first gastral 
sternite, isolates this species within the group. 

Loboponera nasica (Santschi) comb. n. 

(Figs. 8, 11) 

Pachycondyla (Bothroponera) nasica Santschi, 1920: 
6. Holotype worker, GABON: Samkita (F. Faure) 

Worker. TL 4.7-5.6, HL 0.91-0.99, HW 0.86- 
0.95, CI 95-96, SL 0.80-0.90, SI 93-95, AL 1 .82-2.02 
(3 measured). Characters of vigilans group and the 
following. Maximum diameter of eye 0.03-0.04. First 
gastral sternite without a tooth on each side of the 
anteroventral median process but with a median longi- 
tudinal carina. Propodeal declivity with an acute tooth 
near the base; above this the margin has an indistinct 
narrow irregular lamella that may peter out dorsally or 
may have a small dorsal tubercle or blunt tooth. Base of 
first gastral tergite without the bizarre modification of 
basalis (see above). All dorsal surfaces of head and 
body glossy, with longitudinal costulate-rugulose 
sculpture and scattered foveolate punctures. On the 
second gastral tergite coarse longitudinal costae pre- 

This is the most generalised species of the group, 
characterised mainly by its lack of the specialisations 
shown in the other four species. 

Material examined. Ivory Coast: Lamto Field 
Station, nr Toumodi (J. Levieux). Ghana: Tafo (D. 
Leston); Legon (D. Leston); Mampong (P. Room), 
Kade (R. Belshaw). Cameroun: Bakundu (A. Dejean). 
Gabon: Samkita (F. Faure). 

Loboponera subatra sp. n. 

Holotype worker. TL 7.3, HL 1 .32, HW 1 . 1 1 , CI 
84, SL 1.27, SI 1 14, AL 2.38. Characters of vigilans 

group and the following. Maximum diameter of eye 
0.10. First gastral sternite without trace of a median 
longitudinal carina. Anteroventral median process of 
first gastral sternite with an anteriorly directed blunt 
tooth on each side that also arises from the sternite. 
Propodeal declivity with a broad triangular tooth near 
base, the margin above this with a small rounded 
tubercle; dorsum of alitrunk rounds into declivity. 
Short pilosity very dense on all dorsal surfaces, also 
dense on scapes, femora and tibiae. Dorsum of head 
and alitrunk predominantly longitudinally costate- 
rugose, the alitrunk also with foveolae but most of 
these are partially effaced by the longitudinal compo- 
nent; foveolae are most clearly defined on propodeum. 
Second gastal tergite longitudinally costate-rugose, 
the sculpture better defined here than on the first 
tergite, where the foveolate component is more obvious. 

Holotype worker, Cameroun: Nkoemvon, 1980, 
P74 (D. Jackson) (BMNH). 

This relatively large, slender species is recognised 
by the lateral blunt tooth on each side of the 
anteromedian process of the first gastral sternite. In 
vigilans there is usually an angular development in the 
same place, but the latter species has characteristic 
sculpture and has a median carina on the first gastral 
sternite, which is completely absent in subatra. 

Loboponera vigilans sp. n. 

(Fig. 7) 

HOLOTYPE worker. TL 6.3, HL 1 .20, HW 1 . 1 5, CI 
96, SL 1.10, SI 96, AL 2.10. Characters of vigilans 
group and the following. Maximum diameter of rela- 
tively large eye 0. 1 9. First gastral sternite with a median 
longitudinal carina at least on the posterior half of the 
sclerite. First sternite at each side of the anteroventral 
median process forms a short projecting angle or low 
prominence. First gastral tergite in dorsal view with 
bluntly angular anterolateral corners; sides slightly 
constricted behind the corners. Propodeal declivity 
with a blunt tooth near base and another, smaller, tooth 
near apex; the basal teeth appear very broad and coarse 
in dorsal view. Petiole in dorsal view bluntly 
subtriangular, widest posteriorly. All dorsal surfaces of 
head and body with coarse broad foveolate punctures, 
separated by longitudinal costae or elevated narrow 
flat surfaces. The entirety, including bases of foveolae 
and surfaces that separate them, is completely covered 
with extremely fine dense microsculpture and is dull 
and opaque. This microsculpture also occurs on at least 
the upper half of the propodeal declivity and is present, 
though less intensely, on the scapes and legs. 

Paratype workers. TL 6.0-6.6, HL 1 . 16-1 .23, HW 
1.06-1.20, CI 91-98, SL 0.99-1.19, SI 95-103, AL 
1.91-2.14 (10 measured). Maximum diameter of eye 


Holotype worker, Ivory Coast: Abidjan, Banco 
National Forest, 1958, rain forest (E.S. Ross & 
R.E. Leech) (CAS). 

Paratype workers. 4 workers with same data as 
holotype; 1 worker, Ivory Coast: Abidjan, Banco Nat. 
Pk, 14.iii. 1 977, dead wood sample (/. Lobl); 4 workers, 
Tai Forest, 17.x. 1980 (V. Mahnert & J.-L. Perret); 1 
worker, Agboville, Yapo Forest, nr Yapo-Gare, 21- 
22.iii.1977, forest litter (/. Lobl); 6 workers, Nigeria: 
Gambari, 16.vii.1969, rotten log (B. Bolton); 1 worker, 
nrlbadan, IITA, 18. v. 1981, secondary forest, no. 7 (A. 
Russell-Smith); 1 worker, Cameroun: Nkoemvon, 
24.iv.1980 (D. Jackson) (CAS, MCZ, BMNH, SAM). 

Non-paratypic material examined. Cameroun: 
Mvini (A. Dejean); Ottotomo (A. Dejean); Pan Pan (A. 
Dejean). Gabon: La Makande, Foret des Abeilles (S. 
Lewis). Zaire: Kikwit, Kinzambi (A. Dejean). 

Apart from having the largest eyes known in the 
genus, vigilans is quickly identified by its unique 
sculpture. Scanning electron microscope photographs 
of this species appear in Bolton, 1994: 180, figs 493, 


Psalidomyrmex Andre, 1890: 313. Type-species: 
Psalidomyrmexfoveolatus, by monotypy. 

Worker. Ponerine ants with the tribe and group 
characters noted above and also with the following. 

1 . Median portion of clypeus with its anterior margin 
shallowly convex, not forming a projecting lobe. 

2. Labium with upper portion directed anteriorly and 
projecting as a narrow but conspicuous lobe in front 
of the clypeal margin; labral lobe clearly visible in 
full-face view and usually transversely striate. 

3. Mandible triangular to falcate, the apical tooth 
attenuated and the basal angle rounded. 

4. Palp formula 3,4 (dissections of foveolatus, 

5. Posteroventral curve of head in profile without a 

6. Propodeal dorsum with a median longitudinal 
groove or impression. 

7. Mesofemur and metafemur without a mid-dorsal 
longitudinal groove on the basal half. 

QUEEN. Known for feae, foveolatus, procerus: 
slightly larger than the worker, alate, with the above 
characters and the usual modifications associated with 
this caste. 

Male. Known for foveolatus, procerus, 
reichenspergeri: mesotibia and metatibia each with a 
single spur; pretarsal claws with a preapical tooth; 
pygidium truncated, not spinose; notauli absent. For 
general description see Bolton (1975). 


Characters 2 and 3 are autapomorphic; 1 and 5 are 
plesiomorphic. Characters 4 and 6 are possible 
synapomorphies with Plectroctena. The first of these 
(PF 3,4) appears to be the basic maximum count in 
both genera (with subsequent reductions to PF 2,3 and 
PF 2,2 in some Plectroctena species). Character 6 
appears autapomorphic at first glance but queens of 
some species in the Plectroctena minor group show a 
partial (cristata ergatoid) to complete (minor, latinodis 
alates) propodeal groove. In addition, some PI. minor 
workers have a short anterior stub of the groove, so a 
basically synapomorphic state with subsequent loss in 
most Plectroctena must be considered. Character 7 is 
equivocal and is noted under Loboponera comments. 
The genus Psalidomyrmex was revised earlier by 
Bolton (1975). Following the key are taxonomic 
synopses and records of more recently discovered 
material of each species, together with diagnoses of 
species groups within the genus, but formal diagnoses 
of individual species are not repeated as no new spe- 
cies-rank taxa are reported here. 

List of species 

foveolatus group 

foveolatus Andre, 1890 
reichenspergeri Santschi, 1913 

= mandibularis subsp. mabirensis (Arnold, 1954) 
sallyae Bolton, 1975 
procerus group 
feae Menozzi, 1922 

=feae var. impressa Menozzi, 1922 
procerus Emery, 1901 

= longiscapus Santschi, 1920 

= obesus Wheeler, 1922 

= procerus st. collarti Santschi, 1937 
wheeleri Santschi, 1923 

Key to species (workers) 

1 Masticatory margin of mandible concave posterior to the 
elongate apical tooth and with a number of short or 
blunted small teeth at least near the basal angle; overall 
appearance of mandible falcate 2 

- Masticatory margin of mandible straight posterior to the 
elongate apical tooth and edentate throughout its length; 
overall appearance of mandible not falcate 4 

2 Dorsum of first gastral tergite with small punctures, the 
diameters of which are less than the distances separating 
them. Sculpture on anterior half of first gastral tergite 
strongly contrasting to that on anterior half of second 
tergite: the former smooth between the small punctures, 
the latter longitudinally costulate-striate with scattered 
larger punctures. Petiole node in dorsal view longer than 
broad. Large black species, HW > 2.50. (Cameroun. 
Zaire, Uganda) reichenspergeri 


Dorsum of first gastral tergite with coarse foveolate 
punctures, the diameters of which are equal to or greater 
than the distances separating them. Sculpture on anterior 
half of first gastral tergite basically the same as that on 
anterior half of second tergite: either both smooth be- 
tween foveolate punctures, or both longitudinally 
costulate-striate between foveolate punctures. Petiole 
node in dorsal view as broad as or broader than long. 
Smaller red-brown species, HW <2.50 3 

3 Spaces between foveolate punctures on pronotal dorsum 
and first gastral tergite densely striate. Anterodorsal 
margin of mesotibia in its apical half with a row of short 
but strong spine-like traction setae among the normal 
pilosity, the same as the traction setae present at the 
mesotibial apex. Scape relatively short, SI < 75. (Sierra 
Leone, Ivory Coast, Ghana, Nigeria, Cameroun) 

Spaces between foveolate punctures on pronotal dorsum 
and first gastral tergite unsculptured, mostly smooth and 
shining. Anterodorsal margin of mesotibia without spine- 
like traction setae among the normal pilosity; traction 
setae present at the mesotibial apex. Scape relatively 
long, SI > 80. (Ghana) sallyae 

4 Scape relatively longer and head narrower: SI 90- 1 02, CI 
84—89. Petiole node longer that broad in dorsal view. 
(Cameroun, Zaire) wheeleri 

Scape relatively shorter and head broader, SI 79-88, 
CI 91-96. Petiole node broader than long in dorsal view 

5 Frontal lobes smooth and shining over the antennal inser- 
tions. Median portion of clypeus not longitudinally striate. 
(Sao Tome & Principe) feae 

- Frontal lobes uniformly striate over the antennal inser- 
tions. Median portion of clypeus longitudinally striate. 
(Ghana, Cameroun, Gabon, Zaire, Burundi, Uganda) 

Synopsis of species 

foveolatus group 

Mandible falcate; blade of mandible relatively slender, 
narrowing apically and continuing to narrow with- 
out interruption into a much elongated curved apical 

Masticatory margin of mandible concave and with 
weak teeth or crenation on the concave margin distal 
of the basal angle. 

Mandible proximal of rounded basal angle narrowed 
and elongated. 

Psalidomyrmex foveolatus Andre 

Psalidomyrmex foveolatus Andre, 1890: 314. Syntype 
workers, SIERRA LEONE (A. Mocquerys) 


Psalidomyrmex foveolatus Andre; Emery, 1899: 471 
(misidentification); Stitz, 1910: 129; Wheeler, W.M. 
1922: 785; Bernard, 1953: 209; Bolton, 1975: 7. 

Material examined. As Bolton ( 1 975), plus: Ivory 
Coast: Palmeraie Lame (T. Diomande); Iringou (F. 
Krell). Nigeria: Gambari (B. Taylor); Ibadan (A. 
Russell-Smith). Cameroun: Ottotomo (A. Dejean). 

Psalidomyrmex reichenspergeri Santschi 

Psalidomyrmex reichenspergeri Santschi, 1913: 302. 

Holotype worker, CAMEROUN: Molunda (A. 

Reichensperger) (NMB). 
Plectroctena mandibularis subsp. mabirensis Arnold, 

1954: 293, figs 3, 3a. Syntype workers, UGANDA: 

Mabira Forest, 21. v. 1952 (G. Arnold) (SAM, 

BMNH). [Synonymy by Bolton, 1975: 8.] 
Psalidomyrmex mabirensis (Arnold); Bolton, 1974: 

334. [Raised to species and revised combination.] 
Psalidomyrmex reichenspergeri Santschi; Santschi. 

1914b: 288; Wheeler, W.M. 1922: 90, 786; Bolton, 


Material examined. As Bolton ( 1 975 ). 

Psalidomyrmex sallyae Bolton 

Psalidomyrmex sallyae Bolton, 1975: 9, fig. 5. 
Holotype and paratype workers, GHANA: Tafo, 
23.vii.1966, ant ecology sample 120 (D. Lesion); 
paratype worker, GHANA: Tafo, 19.vii.1966, ant 
ecology sample 1 10(BMNH,MCZ.AMNH,SAM). 

Material examined. Known only from type-series. 

procerus group 

Mandible subtriangular; blade of mandible relatively 
broad and with an elongate curved apical tooth. 

Masticatory margin of mandible straight and edentate. 

Mandible proximal of rounded basal angle short and 

Psalidomyrmex feae Menozzi 

Psalidomyrmex feae Menozzi, 1922: 349. Syntype 
workers, queen and male, SAO TOME & PRINCIPE: 
Principe I., Roca Infante Don Henrique, iii.1901, 
100-300 m. (L. Fea) (IE, MCZ). 

Psalidomyrmex feae var. impressa Menozzi, 1922: 
352. Syntype workers and male, SAO TOME & 
PRINCIPE: Principe I., Ro?a Infante Don Henrique, 
ii.1901, 200-300 m. (L. Fea) (IE, MCZ, BMNH). 
[Synonymy by Bolton, 1975: 1 1.] 

Psalidomyrmex feae Menozzi; Bolton, 1975: 11. 

Material examined. Known only from above 


Psalidomyrmex procerus Emery 

Psalidomyrmex procerus Emery, 1901: 50. Syntype 

worker, queens and male, CAMEROUN (Conradt) 

Psalidomyrmex longiscapus Santschi, 1920: 8. 

Holotype queen, GABON: Samkita (F. Faure) 

(NMB). [Synonymy by Bolton, 1975: 12.] 
Psalidomyrmex obesus Wheeler, W.M. 1922: 92, fig. 

19. Syntype workers, ZAIRE: Medje (Lang & 

Chapin) (MCZ, AMNH). [Synonymy by Bolton, 

1975: 12.] 
Psalidomyrmex procerus var. obesus Wheeler; 

Santschi, 1937: 74. [Reduced to variety.] 
Psalidomyrmex procerus st. collarti Santschi, 1937: 

74. Holotype worker, ZAIRE: Ituri, Matenda, 

22.ix.1929 (A. Collart) (MRAC). [Synonymy by 

Bolton, 1975: 12.] 
Psalidomyrmex procerus Emery; Stitz, 1910: 129; 

Wheeler, W.M. 1922: 90, 785 (misidentification); 

Bernard, 1953: 209; Wheeler, G.C. & Wheeler, J. 

1964: 454; Bolton, 1975: 12. 

Scanning electron microscope photographs of this spe- 
cies appear in Bolton, 1994: 175, figs 471, 473. 

Material examined. As Bolton (1975), plus: Cam- 
eroun: Ottotomo (A. Dejean); Abong Mbang (A. 
Dejean). Burundi: Banage (A. Dejean). 

Psalidomyrmex wheeleri Santschi 

Psalidomyrmex wheeleri Santschi, 1923: 263. Syntype 
workers, ZAIRE: Medje, Akenge & Niapu (Lang & 
Chapin) (MCZ, AMNH, MRAC, BMNH, SAM). 

Psalidomyrmex wheeleri Santschi; Bolton, 1975: 13. 

Material examined. As Bolton (1975). 

Dispersal from Psalidomyrmex 

Psalidomyrmex clavicornis Bernard, 1953: 209, fig. 5. 
[Transferred to Pachycondyla, where it is a junior 
synonym of P. talpa (Andre), by Bolton, 1975: 6.] 


Plectroctena F. Smith, 1858: 101. Type-species: 
Plectroctena mandibularis, by monotypy. 

Cacopone Santschi, 1914c: 325. Type-species: 
Cacopone hastifer, by monotypy. [Synonymy by 
Bolton, 1974:313.] 

WORKER. Ponerine ants with the tribe and group 
characters noted above and also with the following. 

1 . Median portion of clypeus with its anterior margin 
shallowly concave. 

2. Clypeal margin anterolaterally with an extensive, 
roughly semicircular, excavation around the man- 
dibular articulation. 


3. Labrum with basal portion not produced into a 
projecting lobe; sclerite minutely but extremely 
densely strongly sculptured. 

4. Mandible stoutly linear, blunt or obliquely trun- 
cated apically, with 0-2 teeth; dorsal surface with a 
longitudinal groove or trench. 

5. Palp formula 3,4 (dissections of cristata, macgeei, 
mandibularis, minor, strigosa), or PF 2,3 
(ugandensis), or PF 2,2 (anops, cryptica, lygaria, 

6. Posteroventral curve of head in profile without a 
projecting flange but anterior portion of occipital 
carina may form a conspicuous crest on the ventral 

7. Propodeal dorsum usually without a median longi- 
tudinal groove but a vestige may be present in some 
individuals of the minor group. 

8. Mesofemur and metafemur with a longitudinal 
(glandular?) groove present mid-dorsally, at least 
on the basal half. 

Queen. Mostly alate but four species have ergatoid 
queens (cristata, dentata, macgeei, mandibularis). All 
have the above characters and alates have the usual 
modifications associated with this caste. Ergatoids 
may be extremely worker-like but usually retain ocelli 
or ocellar vestiges; see Bolton (1974). Queens remain 
unknown in hastifera, laevior, strigosa. 

MALE. Insufficiently known (lygaria, mandibularis, 
minor, subterranea only). For general description see 
Bolton ( 1 974) but this is compromised as lygaria lacks 
the notauli and toothed pretarsal claws that are present 
in the other three. 


Characters 1, 2, 4 and second part of 3 (sculpture) are 
autapomorphic; 6 is plesiomorphic. The possibly 
synapomorphic nature of characters 5 and 7 is discussed 
under Psalidomyrmex. that of 8 under Loboponera. 

The genus Plectroctena was revised earlier by Bol- 
ton (1974). Following the key are taxonomic synopses 
and records of more recently discovered material of 
each species, together with diagnoses of species groups 
within the genus, but formal diagnoses of individual 
species are not repeated as no new species-rank taxa 
are reported here. 

List of species 

hastifera group 
anops Bolton, 1974 
hastifera (Santschi, 1914) 
minor group 

cristata Emery, 1 899 

= cristata var. semileavis Santschi, 1 924 
dentata Santschi, 1912 
= emeryi Santschi, 1924 


latinodis Santschi, 1924 

minor Emery, 1 892 

= gabonensis Santschi, 1919 syn. n. 
= minor var. perusta Santschi, 1924 
= minor var. liberiana Santschi, 1924 
= minor var. insularis Santschi, 1924 
mandibularis group 

cryptica Bolton, 1974 

gestroi Menozzi, 1922 

laevior Stitz, in Santschi, 1924 

lygaria Bolton, Gotwald & Leroux, 1979 

macgeei Bolton, 1974 

mandibularis F. Smith, 1858 

= caffra Spinola, 1851 (nomen nudum) 
= caffra st. major Forel, 1 894 
= minor st. conjugata Santschi, 1914 
= mandibularis st. integra Santschi, 1924 

[mandibularis st. strigosa var. strialiventris Stitz, in 

Santschi, 1924; unavailable name] 

strigosa Emery, 1 899 

subterranea Arnold, 1915 
=punctatus Santschi, 1924 

ugandensis Menozzi, 1933 

Key to species (workers) 

1 Propodeum with a continuous cuticular lamella that forms 
an uninterrupted projecting flange across the dorsum and 
down the sides of the declivity. Apical half of each 
mandible swollen, basal tooth always absent. Head 
relatively narrow, CI < 80 2 

Propodeum either without a projecting cuticular lamella 
or with a lamella that is confined to each side of the 
declivity and does not extend across the dorsum. Apical 
half of each mandible not swollen, basal tooth usually 
present. Head relatively broad, CI > 85 3 

2 Dorsal surfaces of head, alitrunk. petiole and first and 
second gastral tergites with numerous short erect to 
suberect hairs. Eyes present, minute. (Ghana) 


- Dorsal surfaces of head, alitrunk. petiole and first and 
second gastral tergites without standing hairs of any 
form. Eyes absent. (Ghana) anops 

3 Dorsum of first gastral tergite with an anteriorly located 
transverse groove or impression which may extend the 
width of the tergite or be confined to the median quarter 
of the width of the sclerite 4 

Dorsum of first gastral tergite without trace of an anteriorly 
located transverse groove or impression 7 

4 Ventral surfaces of the head and usually also the sides of 
the head, at least below and behind the eyes, with the 
spaces between punctures finely striate. Funicular 
segments 3—4 as long as, or longer than broad 5 

Ventral surfaces of the head and the sides of the head 
below and behind the eyes with the spaces between 


punctures smooth and shining. Funicular segments 3-4 
broader than long, usually markedly so 6 

5 Very large species with relatively very long mandibles, 
HL > 4.0, Ml > 90. Transverse groove on first gastral 
tergite strongly developed across the width and preceded 
by a thick crest of cuticle; very conspicuous in profile. 
(Cameroun, Zaire, Uganda, Kenya) cristata 

Smaller species with relatively shorter mandibles. HL < 
4.0, MI < 90. Transverse groove on first gastral tergite 
faint and poorly developed, only visible in the middle of 
the tergite and not preceded by a thick crest of cuticle; 
scarcely discernible in profile. (Republic of Congo, Za- 
ire, Angola. Uganda) dentata 

6 Petiole node in dorsal view as broad as long. (Cameroun, 
Zaire) latinodis 

- Petiole node in dorsal view longer than broad. (Sierra 
Leone, Liberia. Ivory Coast. Ghana, Togo, Nigeria, Cam- 
eroun. Equatorial Guinea. Gabon. Zaire) minor 

7 Dorsum of head coarsely sculptured with foveolae or 
large pits whose diameters are greater than the distances 
that separate them or which are adjacent. Mesonotum and 
propodeum with a polished, virtually unsculptured. 
median longitudinal strip. (Liberia. Ivory Coast, Ghana) 

Dorsum of head finely sculptured with small punctures 
whose diameters are less than the distances that separate 
them. Mesonotum and propodeum without a polished, 
virtually unsculptured, median longitudinal strip 8 

8 Labrum shallowly transversely concave and with a sharply 
incised median longitudinal groove that extends the en- 
tire length of the sclerite and completely breaks the fine 
surface sculpture. Mandible edentate. (Ghana, Nigeria) 


- Labrum without a median longitudinal groove, the fine 
surface sculpture uninterrupted across the entire shallowly 
transversely concave sclerite. Mandible at least with a 
strong basal tooth; usually also with a second, smaller, 
tooth in the apical half 9 

9 Dorsal surfaces of head and alitrunk with numerous short 
erect or suberect hairs present. (Principe I.) gestroi 

Dorsal surfaces of head and alitrunk without standing 
hairs 10 

1 Ventral surfaces of head without striation between the 
punctures 1 1 

- Ventral surfaces of head with striation between the punc- 
tures, at least anteriorly 14 

1 1 Propodeal declivity armed near its base with a stout 
triangular tooth; lamella not developed dorsal to the 
tooth. Larger species. HL > 3.0; full adult colour blackish 
brown to black. (Tanzania) laevior 

- Propodeal declivity not armed near its base with a stout 
triangular tooth; lamella fully developed and extends 
almost the height of the declivity. Smaller species, HL < 
2.5; full adult colour orange-brown or dark red 12 



12 Foveolate punctures on dorsum of head each with an 
appressed small hair arising from its centre, the hair 
directed toward the cephalic midline. (Ivory Coast) 


- Foveolate punctures on dorsum of head without appressed 
small hairs arising from their centres 13 

1 3 Basal tooth on mandible with distal (anterior) and proxi- 
mal (posterior) surfaces that are of about equal length and 
slope; proximal base of tooth linked to mandible base by 
a shallow low ridge. Colour orange-brown. (Kenya, 
Malawi, Zimbabwe, Namibia) subterranea 

Basal tooth on mandible with vertical distal (anterior) 
surface but without a defined proximal (posterior) surface; 
instead the apex of the tooth is extended to the mandible 
base as an oblique high ridge. Colour dark red. (Ivory 
Coast, Cameroun, Zaire, Uganda) ugandensis 

14 Leading edge of scape with a row of suberect to erect 
short hairs. All dorsal surfaces of head and body entirely 
covered with extremely dense fine striate sculpture. 
(Kenya, Tanzania, South Africa) strigosa 

Leading edge of scape without suberect to erect short 
hairs. Sculpture variable in density and intensity but not 
all dorsal surfaces of head and body entirely covered with 
extremely dense fine striate sculpture. (Ethiopia, Kenya. 
Uganda, Burundi, Zaire, Angola, Tanzania, Malawi, 
Mozambique. Zimbabwe, Botswana, South Africa) 

Synopsis of species 

hastifera group 

Propodeal lamellae form a continuous flange or rim 
around the sides and dorsum of the declivity. 

Mandible with basal tooth absent. 

First gastral tergite without an anteriorly situated trans- 
verse groove or impression on the dorsum. 

Petiole node in profile long and low. 

Head considerably longer than broad, CI < 80. 

Plectroctena anops Bolton 

Plectroctena anops Bolton, 1974: 319, figs 4, 7. 
Holotype worker, GHANA: Tafo, 8.ix.l966, ant 
ecology sample 249c (D. Leston) (BMNH). 

Material examined. As Bolton (1974), plus: 
Ghana: Kwadaso (J. Plisko). 

Plectroctena hastifera (Santschi) 

Cacopone hastifer Santschi, 1914c: 325, fig. 11. 

Holotype worker, GHANA: Aburi (F Silvestre) 

Plectroctena hastifera (Santschi); Bolton, 1974: 320. 

[Revised combination.] 

Material examined. As Bolton (1974). 

minor group 

Propodeal lamellae restricted to sides of the declivity, 
often very weakly developed. 

Mandible with basal tooth present and usually also 
with a second, smaller, more apically situated tooth. 

First gastral tergite with an anteriorly situated trans- 
verse groove or impression on the dorsum. 

Petiole node in profile as high as or higher than long. 

Head relatively broad, CI 89-97. 

Plectroctena cristata Emery 

Plectroctena cristata Emery, 1 899: 470. Syntype work- 
ers, CAMEROUN (Conradt) (MCSN). 

Plectroctena cristata var. semileavis Santschi, 1924: 
163 (variant spelling as semilaeve: 173). Holotype 
worker, ZAIRE: Luebo, Kamaiembi, 22. ix. 1 92 1 (H. 
Schouteden) (MR AC). [Synonymy by Bolton, 1974: 

Plectroctena c rista ta Emery; Stitz, 1910: 129; Wheeler, 
W.M. 1922: 88, 783; Santschi, 1924: 163; Bolton, 

Material examined. As Bolton (1974), plus: 
Uganda: Busongoro(G.D.//. Carpenter). Cameroun: 
Tissongo (D. Jackson). 

Plectroctena dentata Santschi 

Plectroctena minor var. dentata Santschi, 1912: 150. 
Syntype workers, ANGOLA: Benguela, Cucala (J. 
Crochet) (NMB, MRAC). 

Plectroctena dentata Santschi; Santschi, 1924: 164. 
[Raised to species.] 

Plectroctena emeryi Santschi, 1924: 164. Holotype 
queen (ergatoid, not worker), REPUBLIC OF 
CONGO (J. de Gaule) (NMB). [Synonymy by Bol- 
ton, 1974: 322.] 

Plectroctena dentata Santschi; Bolton, 1974: 322. 

Material examined. As Bolton (1974). 

Plectroctena latinodis Santschi 

Plectroctena latinodis Santschi, 1924: 165, fig. 2a. 

Syntype worker and queen: ZAIRE: Congo da 

Lemba (R. Mayne) (NMB, MRAC). 
Plectroctena latinodis Santschi; Bolton, 1974: 323. 

Material examined. As Bolton (1974). 

Plectroctena minor Emery 

Plectroctena minor Emery, 1892: 556, pi. 15, figs 1,2. 

Holotype queen, IVORY COAST: Assinie, vii- 

viii.1886 (C. Alluaud) (MCSN). 
Plectroctena gabonensis Santschi, 1 9 1 9a: 336. Syntype 

workers, GABON: Libreville, l.xii.1897 (Chalot) 

and GABON: Samkita, 1914 (F. Fame) (NMB). 

Syn. n. 



Plectroctena subterranea st. gabonensis Santschi, 
1919b: 90. [Second description as new, based on 
same specimens as above.] 

Myopias subterranea subsp. gabonensis (Santschi); 
Wheeler, W.M. 1922: 785. [Revised combination.] 

Plectroctena gabonensis Santschi; Santschi, 1924: 170. 
[Revived combination and revived status.] 

Plectroctena minor var. perusta Santschi, 1924: 168, 
fig. 2b. Syntype workers, CAMEROUN: 
Barumbistation (Preuss) (NMB). [Synonymy by 
Bolton, 1974:324.] 

Plectroctena minor var. liberiana Santschi, 1924: 169, 
fig. 2c. Holotype worker, LIBERIA (NMB). [Syn- 
onymy by Bolton, 1974: 324.] 

Plectroctena minor var. insularis Santschi, 1924: 169, 
fig. 3a. Holotype worker, EQUATORIAL GUINEA: 
Fernando Po I. (Conradt) (MCSN). [Synonymy by 
Bolton, 1974:324.] 

Plectroctena gabonensis Santschi; Wheeler, W.M. 
1922: 783; Bolton, 1974:323. 

Plectroctena minor Emery; Emery, 1 902: 32; Wheeler, 
W.M. 1922: 88, 784; Santschi, 1924: 167: Bolton, 


Accretion of new material has gradually eliminated the 
supposed differences of size, colour, relative size of 
eye and minor structural features that were formerly 
invoked to separate minor and gabonensis (Santschi, 
1924; Bolton, 1974); the two are newly synonymised 

Scanning electron microscope photographs of this 
species appear in Bolton, 1994: 175, figs 470, 472. 

Material examined. As Bolton (1974), plus: 
Ghana: Tafo (D. Leston); Wiawso (D. Leston); Odomi 
Riv. (D. Leston). Togo: Palime, Klouto (Vit). Camer- 
oun: Ottotomo (A. Dejean); Ndupe (A. Dejean); Nzi 
(A. Dejean); Nkoemvon (D. Jackson). Gabon: La 
Makande, Foret des Abeilles (S. Lewis). Zaire: 
Kinzambi (A. Dejean). 

mandibularis group 

Propodeal lamellae restricted to sides of the declivity, 
often very weakly developed. 

Mandible with basal tooth usually present (absent only 
in macgeei); usually also with a second, smaller, 
more apically situated tooth. 

First gastral tergite without an anteriorly situated trans- 
verse groove or impression on the dorsum. 

Petiole node in profile usually as high as or higher than 
long (relatiely long and low in macgeei). 

Head relatively broad, CI 86-95. 

Plectroctena cryptica Bolton 

Plectroctena cryptica Bolton, 1974: 327, fig. 5. 
Holotype worker, GHANA: Tafo, 2. i. 1969, on mud 

below dam (B. Bolton) (BMNH). Paratype workers, 
IVORY COAST: Lamto (Toumodi), ll.iv.1968, 
sample AA279N2; Lamto., sample 
AA334N4; and Lamto, 21. ii. 1969 (J. Levieux) 

Material examined. As Bolton (1974), plus: Ivory 
Coast: Tai Forest (T. Diomande); Abidjan, 
Adiopodoume (/. Lobl). Ghana: Mt Atewa(D. Leston). 

Plectroctena gestroi Menozzi 

Plectroctena gestroi Menozzi, 1922: 348, fig. 1. 
Syntype workers and queen, SAO TOME & 
PRINCIPE: Principe I.: Roca Infante Don Enrique, 
hi. 1900 (Z.. Fea)(\E. MCZ). 

Plectroctena gestroi Menozzi; Bolton, 1974: 328. 

Material examined. Known only from type-series. 

Plectroctena laevior Stitz 

PU-( iroctena mandibularis st. laeviorSutz, in Santschi, 
1924: 163, fig. Id. Holotype worker, TANZANIA: 
Kiwugebeit (Kadt) (MNHU). 

Plectroctena laevior Stitz; Bolton, 1974: 329. [Raised 
to species.] 

Material examined. Known only from holotype. 

Plectroctena lygaria Bolton, Gotwald & 

Plectroctena lygaria Bolton. Gotwald & Leroux, 1 979: 
373, figs 1, 2. Holotype worker, paratype workers, 
queens and males, IVORY COAST: Lamto, 1974. foret galerie du Bandanna ( W.H. Gotwald 
& J.M. Leroux) (BMNH, MCZ, MNHN, NMB, 

Material examined. Known only from type-series. 

Plectroctena macgeei Bolton 

Plectroctena macgeei Bolton, 1974: 330, figs 1, 8. 
Holotype worker, NIGERIA: Western State, 
Gambari, 28.x. 1969, amongst termites under log (B. 
Bolton) (BMNH). 

Material examined. As Bolton (1974), plus: 
Ghana: Tafo (D. Leston); Ashanti, Juaso {R. Belshaw); 
Atewa For. Res., nr Kibi (R. Belshaw). Nigeria: 
Gambari (B. Taylor); Ibadan (B.R. Critchley). 

Plectroctena mandibularis F. Smith 

Plectroctena mandibularis F. Smith, 1858: 101, pi. 7, 
figs 1-5. Syntype queen (ergatoid, not worker) and 
male, SOUTH AFRICA: Natal, Durban (= Port 
Natal) (Gueinzius) (BMNH). 

Ponera caffra Spinola, 1851: 53; Spinola, 1853: 69. 



Nomen nudum (attributed to Klug). [Spinola mate- 
rial referred to mandibularis by Roger, 1861 : 41.] 

Plectroctena caffra (Klug); Roger, 1861:41 [combina- 
tion]; Emery, 1892: 556 [as valid species, in error]; 
Dalla Torre, 1893: 31 [as senior synonym of 
mandibularis, in error]. 

Plectroctena caffra r. major Forel, 1894: 74. Holotype 
queen (ergatoid, not worker), MOZAMBIQUE: 
Delagoa (P. Berthoud) (MHN). [Synonymy by Em- 
ery, 1899: 469; Santschi, 1924: 160; Bolton, 1974: 

Plectroctena mandibularis var. major Forel; Emery, 
191 1:95; Santschi, 1914a: 54; Arnold, 1915:86. 

Plectroctena minor st. conjugata Santschi, 1914d: 8. 
Syntype workers and queen: SOUTH AFRICA: 
Natal, Stamford Hill, Charlestown, 30.iv.1905, and 
SOUTH AFRICA: Zululand (/. Trdgardh) (NMB, 
MCZ, MRAC). [Synonymy by Arnold, 1926: 209; 
revived from synonymy by Bolton, 1974: 326; syn- 
onymy reaffirmed by Villet, McKitterick & 
Robertson, 1999: 282.] 

Plectroctena mandibularis var. Integra Santschi, 1 924: 
161. Syntype worker, KENYA: Nairobi, Wa Kikongo 
et Masai, 1904 (C. Alluaud); syntype male, KENYA: 
Bura, Wa Taita, 1904 (C. Alluaud (NMB). [Syn- 
onymy by Bolton, 1974: 330.] 

[Plectroctena mandibularis st. strigosa var. 
strialiventris Stitz, in Santschi, 1924: 162 (with 
variant spellings strativentris and striativentris: 1 62). 
Unavailable name; material (from Malawi) referred 
to mandibularis by Bolton, 1974: 330.] 

Plectroctena conjugata Santschi; Wheeler, W.M. 1 922: 
785; Santschi, 1924: 166 [raised to species]; Bolton, 
1974: 326; Wheeler, G.C. & Wheeler, J. 1989: 52. 

Plectroctena mandibularis F Smith; Gerstacker, 1 873: 
346; Emery, 1899: 469; Forel, 1913: 108; Wheeler, 
W.M. 1922: 783; Santschi, 1924: 160; Bolton, 1974: 
330; Villet, McKitterick & Robertson, 1999: 282. 

Material examined. As Bolton (1974), plus: 
Kenya: Shimba Hills (B. Holldobler); Kajiado (G 
Nyamasyo); Olikoriti, nr Kajiado (M.G. Lepage). 
Burundi: Bujumbura (A. Dejean). Tanzania: Old 
Shinyanga (O.W. Richards): Mkomazi Game Res., 
Ibanya (A. Russell-Smith). Botswana: Maxwee (A. 
Russell-Smith). South Africa: Transvaal, Hoedspruit 
(C. Peeters); Natal, Mkuzi Res. (C. Peeters); Cape 
Prov., Grahamstown (F. Jacot-Guillarmod); 
Grahamstown (W.L. Brown); Grahamstown (L.S. 
Naylor); Cape Prov., nr Pt Alfred (H.G. Robertson); 
Cape Prov., Alexandria (H.G. Robertson). 

Plectroctena strigosa Emery 

Plectroctena mandibularis var. strigosa Emery, 1899: 
469. Holotype worker, SOUTH AFRICA: Natal 
(Staudinger & Bang-Haas) (MCSN). 

Plectroctena mandibularis var. strigosa Emery ; Arnold, 

1915: 88; Wheeler, W.M. 1922: 784. 
Plectroctena cristata st. strigosa Emery; Santschi. 

1924: 161. 
Plectroctena strigosa Emery; Bolton, 1974: 332. 

[Raised to species.] 


Hamish Robertson (SAM) has drawn attention to the 
fact that material from Kenya and Tanzania that is 
referred to strigosa fits the original description, but no 
specimen matching the Kenyan/Tanzanian material 
has yet been seen from Natal, the type-locality. It is 
possible that the name is misapplied or the specimen 
mislabeled (Emery mentions a specimen from Zanzi- 
bar in his discussion). A reappraisal of the holotype 
will be necessary to resolve the situation. 

Material examined. As Bolton (1974), plus: 
Kenya: Tiwi Beach (C.J. Powles); Shimba Hills (B. 
Holldobler); Shimba Hills (C. Peeters); Gedi Natl. 
Monument (PS. Ward). Tanzania: Mkomazi Game 
Res., Ibaya (A. Russell-Smith). 

Plectroctena subterranea Arnold 

Plectroctena subterranea Arnold, 1915: 84, pi. 3, figs 

23, 23a. Syntype workers and queen, ZIMBABWE: 

Bulawayo, (G. Arnold), and Shiloh (G 

Arnold) (BMNH, SAM). 
Myopias subterranea (Arnold); Wheeler, W.M. 1922: 

785. [Revised combination.] 
Plectroctena subterranea Arnold; Santschi, 1924: 171. 

[Revived combination.] 
Plectroctena punctatus Santschi, 1924: 170. Holotype 

male, KENYA: Bura Wa Taita, hi. 19 12, 1050 m, st. 

6 1 (C. Alluaud & R. Jeannel) (NMB) [Synonymy by 

Bolton, 1974:333.] 

Material examined. As Bolton (1974), plus: 
Kenya: Kibwezi (S.A. Neave). [Also recorded from 
Namibia by Robertson (2000).] 

Plectroctena ugandensis Menozzi 

Plectroctena ugandensis Menozzi, 1933: 99, fig. 2. 
Holotype queen, UGANDA: Bussu (E. Bayon) (lo- 
cation of holotype not known, presumed lost). 

Plectroctena ugandensis Menozzi; Bolton, 1974: 334. 

Material examined. As Bolton ( 1 974), plus: Ivory 
Coast: Tai Forest (T. Diomande). Cameroun: nr 

Yaounde (G Perron); Nzi (A. Dejean). 

Disperal from Plectroctena 

Plectroctena mandibularis subsp. mabirensis Arnold, 
1954: 293. [Transferred to Psalidomyrmex by Bol- 
ton, 1974: 334; junior synonymy with P. 
reichenspergeri by Bolton, 1975: 6 (see above).] 




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Willey, R.B. & Brown, W.L. Jr. 1983. New species of the ant genus 

Myopias. Psyche 90: 249-285. 



Figs. 1-5. Loboponera workers: 1-3, heads in full-face view of 1. basalis; 2. politula: 3. obeliscata: 4-5, body profiles of 4, 
obeliscata; 5, politula. 



Figs. 6-11. Loboponera workers: 6-8, body profiles of 6, trica; 7, vigilans; 8, nasica; 9, profile of petiole and base of gaster 
of basalis; 10-11, dorsal view of petiole and gaster of 10, basalts; 1 1, nasica. 


Bull. nat. Hist. Mus. Land. (Ent.) 71( 1 ): 19-58 Issued 27 June 2002 

New Neotropical Nepticulidae 
(Lepidoptera) from the western 
Amazonian rainforest and the 
Andes of Ecuador 


Department of Zoology, Vilnius Pedagogical University, 39. Studentu str, Vilnius 
2004 LT, Lithuania 


Department of Zoology, Vilnius Pedagogical University, 39, Studentu str, Vilnius 
2004 LT, Lithuania 


Department of Entomology, The Natural History Museum, Cromwell Road, London 
SW7 5BD, UK 


Synopsis 19 

Introduction 19 

Material and methods 20 

Abbreviations of institutions 20 

Acknowledgements 21 

Descriptions of new species 21 

Enteucha Meyrick 21 

Stigmella Schrank 23 

Fomoria Beime 26 

Acalyptris Meyrick 27 

References 34 

Index 34 

SYNOPSIS. Fieldwork in 2000-2001 in the upper Amazon basin and the Andes (Ecuador) has 
yielded 16 new species of Nepticulidae, described here. Four genera are represented: Enteucha 
Meyrick (two species: E. acuta. E. guajavae), Stigmella Schrank ( lour species: S. montanotropica, 
S. nubimontana, S. rubeta, S. austroamericand), Fomoria Beime (two species: F. repanda, F. 
tabulosa) and Acalyptris Meyrick (8 species: A. ecuadoriana, A. onorei, A. basihastatus, A, 
pseudohastatus, A. articulosus, A. rotundas. A. amazonius, A. insolentis); the authors of all new 
taxa are Puplesis & Diskus. Larvae have been reared from three genera of different plant families: 
Psidium CMyrtaceae), Acalypha (Euphorbiaceae) and Rubus (Rosaceae). The adult moths of all 1 6 
species treated are illustrated in black and white, with line drawings of genitalia and. for four 
species, drawings of leaf-mines. 


Description of the Neotropical fauna began in the late 
1 9th and early 20th century with descriptions of species 

by Zeller (1877) and Meyrick (1915, 1931). Further 
Nepticulidae are a very specialized, isolated family of species were described by Forbes & Leonard (1930), 
primitive monotrysian Microlepidoptera with a world- Bourquin ( 1 962), Davis ( 1 978, 1 984, 1 985), Wilkinson 
wide distribution and about 750 described species. (1981) and Newton & Wilkinson (1982). A consider- 

© The Natural History Museum. 2002 



able boost was given to the number of species known 
from the Neotropical region by Puplesis & Robinson 
(2000) who described as new 33 species, mostly from 

In this paper we report upon the results of field- 
collecting by Rimantas Puplesis (VPU) and Simon R. 
Hill (UW) in 2000-2001 in the upper Amazon basin 
and the Andes of Ecuador, areas unrepresented among 
material reviewed by Puplesis & Robinson (2000). 
From these new collections we describe below 16 new 
species - a significant addition to our knowledge of the 
Amazonian and Andean nepticulids - and provide 
larval rearing data from three host genera in different 
plant families. 

This ongoing collaborative project (for previous 
publications, see Puplesis & Robinson, 1999; 2000) 
was undertaken at the Natural History Museum, Lon- 
don with the support of the Royal Society (London) 
and Professor Hering Memorial Fund. It combines 
Robinson's interests in Microlepidoptera biodiversity 
with Puplesis's and Diskus's expertise and interest in 
inventorying the families of primitive 
Microlepidoptera. We hope that further documenta- 
tion of the Neotropical fauna and illustration of newly 
discovered species will stimulate further studies of the 
Nepticulidae of South America; we are very conscious 
that there is probably a great deal more yet to be 
discovered and described. 


Three primary collecting-sites were utilized in the 
upper Amazon Basin, in addition to brief collecting 
trips within the Ecuadorian Andes, covering the west- 
ern and eastern slopes and altitudes from 700 m to 
4200 m. Primary sites were: the Yasuni Research Station 
(Napo Region, SE of Coca, rainforest at 260 m, 
76°36'W, 00°38'S) and Jatun Sacha Biological Station 
and Misahualli (both localities in Napo Region, SE of 
Tena, premontane tropical forest at 400-500 m, 
77°36'W, 01°04'S). 

Two main collecting methods were used: mining 
larvae were collected and reared, and adults were 
collected at light. Mined leaves (or other plant parts) 
were placed in Petri dishes which were then checked 
regularly for emerged adults. Emergence occurred 
within 1-3 weeks. Adult moths were collected by 
attracting them to mercury-vapour light from a lamp 
suspended slightly above eye-level and 5-10 cm in 
front of a white screen, rather closer than is usual in the 
standard method for light-collecting (described by 
Robinson etal., 1994) in which the lamp is about 0.5 m 
from the illuminated surface. A portable Honda EX 
350 generator was used as a power-source. As many 
different habitats and sites as possible were sampled. 

Moths attracted to the screen were collected into small 
glass tubes and pinned after killing with ethyl acetate 

Genitalia were prepared following the method 
described by Robinson (1976). After maceration of the 
abdomen in 10% KOH and subsequent cleaning, male 
genital capsules were removed from the abdomen and 
mounted ventral side uppermost. In cases of compli- 
cated armature, the genitalia were studied and sketched 
in glycerin before fixation. The aedeagus was removed 
from and mounted alongside the genital armature. 
Female genitalia were removed entirely from the 
abdomen, cleaned and mounted ventral side upper- 
most. Genitalia and abdominal pelts of both sexes were 
stained with Chlorazol Black (Direct Black 38/Azo 
Black) and mounted in Euparal. 

Forewing length is expressed as a range, where 
availability of material made this possible, measured 
along the costa from the wing base to the apex of the 
cilia. Wingspan was measured from the tip of the left 
wing to the tip of the right wing, where well-mounted 
specimens were available; in other cases the forewing 
length was doubled and the thorax width added. 

Illustrations of the adults and leaf-mines were made 
in Indian ink by Mr Arturas Skorupskas (VPU) using 
preliminary sketches and notes by R. Puplesis and with 
additional observations using a stereoscopic micro- 
scope (MBS- 10). Adults are illustrated at the same 
scale to indicate comparative size. It should be noted 
that the illustrations of adults are idealized and gener- 
alized. Unfortunately black and white drawings cannot 
show the metallic lustre and coloured iridescence char- 
acteristic of most Nepticulidae; details of such colours 
have been incorporated into species descriptions. 

Genitalia and wing venation drawings were made 
by Puplesis using a camera lucida, mainly from perma- 
nent slides, but occasionally from temporary glycerin 

Depositories of types are given in the species 
descriptions: either BMNH - The Natural History 
Museum, London, UK (formerly British Museum 
(Natural History)) or VPU - Vilnius Pedagogical 
University, Vilnius, Lithuania. 


BMNH The Natural History Museum, London, UK 

(formerly British Museum (Natural History)) 
INEFAN Instituto Ecuatoriano Forestal de Areas Naturales 

y Vida Silvestre, Ecuador 
NNM Nationaal Natuurhistorisch Museum (Naturalis), 

Leiden, Netherlands 
PUCE Pontificia Universidad Catolica del Ecuador, 

Quito. Ecuador 
VPU Vilnius Pedagogical University, Vilnius, 



USNM National Museum of Natural History, Washing- 
ton DC, USA (formerly United States National 
UW University of Westminster, London, UK 

ZMUC Zoologisk Museum, University of Copenhagen, 
Copenhagen, Denmark 

Acknowledgements. Mr Ole Karsholt and Prof. Niels P. 
Kristensen (ZMUC) provided the initial stimulus for the 
present project together with generous support during its 

Mr Simon R. Hill (UW) is thanked for his selfless and 
generous help with planning fieldwork. transport of equip- 
ment, technical assistance and companionship during the 
Ecuador collecting-trips in 2000 and 2001 . 

We are extremely grateful to Dr Giovanni Onore (PUCE) 
for all necessary preparations for a study permit to collect 
material in Ecuador and for scientific collaboration. Special 
thanks are due to Miss Ruth Boada and Miss Palma Lucy 
Elizabeth Baldeon (PUCE ) for their generous help and friend- 
liness during Puplesis's and Hill's fieldwork in Ecuador. We 
thank botanists Jaime Jaramillo A. and David Padilla Z. 
(PUCE) for the identification of numerous host-plant samples 
collected in Ecuador. Our very special thanks extend to bota- 
nist Alvaro Javier Perez (PUCE), who kindly, efficiently and 
very professionally assisted us during the last stage of this 

We are grateful to Arturas Skorupskas (VPU) for making 
Indian-ink drawings of the adults and the mines. 

The first author is grateful to Kevin R. Tuck (BMNH) for 
generous support during the course of this work. 

This project was partially conducted during Puplesis's 
study visits to BMNH under the Royal Society/Lithuanian 
Academy of Sciences Exchange Programme in 2000 and 
2001. We are most grateful to the Royal Society of London 
and Professor Hering Memorial Fund (administered by the 
British Entomological and Natural History Society) for sup- 
port. We are very grateful to INEFAN (Ecuador) for a study 
permit to collect in Ecuador. Puplesis thanks the Trustees and 
the Keeper of Entomology at The Natural History Museum. 
London, for study facilities and access to collections. We 
thank Kevin Tuck, Malcolm Scoble and an anonymous re- 
viewer for refereeing this paper, and for their perceptive and 


For generic diagnoses and synonymy, see Puplesis & 
Robinson (2000). 


Enteucha acuta Puplesis & Diskus sp. n. 

(Figs 4, 26-28) 


Male (Fig. 4). Forewing length: about 1 .6 mm. Wing- 
span: about 3.4-3.5 mm. Head: palpi brownish cream; 
frontal tuft ochreous [heavily rubbed in the single 
available example]; collar probably cream [totally 
rubbed off in the available specimen]; eyecaps cream, 
large; antenna brown, comprising of about 17-18 seg- 
ments. Thorax, tegulae and forewing brown; area of 
forewing before and after fascia dark brown; fascia 
postmedial, shining silver, very distinctive, perpen- 
dicular to forewing. Cilia grey. Underside of forewing 
brown or brownish. Hindwing lanceolate, brownish 
grey; its cilia concolorous. No androconia on forewing 
or hindwing. Legs brownish cream, darkened with 
fuscous brown on upperside. Abdomen brown on 
upperside, paler on underside. 

Female. Unknown. 

Genitalia d (Figs 26-28). Capsule 230-233 m long. 
Tegumen band-like, laterally slightly extended into 
papillated sublateral lobes. Uncus triangular with 
single, well-sclerotized caudal papilla. Gnathos with 
large posterior process and triangular lateral arms; 
central plate weakly developed, in shape of narrow 
band; anterior processes apparently absent or as tiny 
anterior papillae only. Val va 1 60- 1 70 urn long; slender, 
curved inwardly, with broader basal one-quarter to 
one-third, and with pointed apex. Transtilla apparently 
absent (i.e., no sclerotized transverse bar developed), 
however basal parts of valva joined ventrally. Juxta 
present, a very large, medium or weakly sclerotized 
lobe reaching gnathos, basally fused with valva. Vin- 
culum relatively short, constricted and smoothly 
rounded distally ; no anterior excavation or lateral lobes. 
Aedeagus 165-170 urn long, with one very large 
cornutus and two smaller ones one-quarter or less 
length of major cornutus; apex of aedeagus with small 
sharp denticles and long, slender, cornuti-like 
sclerotized wrinkles. 

Biology. Single adult collected in January. 

Diagnosis. Distinctive species; among all currently 
known species of the genus strongly differing by the 
presence of a large cornutus. In external features, 
particularly the fascia, it bears a superficial resem- 
blance to the Neotropical gilvafascia (Davis) and 
snaddoni Puplesis & Robinson; however the fascia of 
the new species is shining silver, and there is no white 
apical spot on the forewing. 


Lowland Amazon rainforest 

Condition of type material. The single speci- 
men is badly pinned, the head is rubbed, however the 
wings are well preserved with clear patterning. 

Material examined. Holotype <3 : Ecuador: Napo 
Region, SE of Coca, near Rio Tiputini,Yasuni National 



Park, 260 m, 15-25.i.2000 (Puplesis & Hill) genitalia 
slide no. 29604 (BMNH). 

Remarks. This remarkable species represents the 
first member of the genus to be found in Amazonia, and 
it is described from only a single specimen because of 
the very distinct characters of the genitalia, which 
exhibit considerable apomorphic modification in com- 
parison with other related species. 

Enteucha guajavae Puplesis & Diskus sp. n. 

(Figs 2, 3, 20-25, 89, 90, 100, 101) 

MALE (Fig. 3). Forewing length: 1.4- 1.6 mm. Wing- 
span: 3.1-3.6 mm. Head: palpi brownish cream; frontal 
tuft ochreous orange; collar cream and paired; eyecaps 
cream and glossy; antenna fuscous on upperside and 
grey on underside, comprising of about 21-24 
segments. Thorax and tegulae fuscous with some purple 
iridescence; forewing of same colour, with two striking 
shiny silvery fasciae: basal fascia almost half width of 
postmedial. Cilia greyish with distinct cilia-line. 
Underside of forewing fuscous. Venation with Sc, four 
radials, one medial, but no cubital vein (Fig. 2). 
Hindwing lanceolate, covered with brownish cream 
androconial scales (especially dense on underside of 
wing, making the hindwing the same colour as the 
underside of the abdomen); cilia of hindwing grey. 
Legs brownish cream, darkened with fuscous brown 
on upperside. Abdomen fuscous on upperside but ochre- 
ous cream on underside. 

Female. Hindwing grey. Otherwise as in male. 

Genitalia d (Figs 20-25). Capsule 317-285 pm 
long. Tegumen band-like, distally widely rounded. 
Uncus with two short but broad papillated lateral lobes. 
Gnathos in shape of inverted V; caudally directed part 
weakly sclerotized and rounded; lateral arms very 
slender. Valva 140-155 pm long; slender, curved 
inwardly, with broader and strongly sclerotized basal 
one-quarter to one-third and with pointed apex. Distinct 
transtilla apparently absent; however, a weakly 
sclerotized transverse bar is evident between basal 
processes of valvae; valvae additionally connected 
ventrally by ventral sclerite which is fused with vincu- 
lum plate. Juxta weakly developed and without clearly 
defined shape. Vinculum very long and widely rounded 
distally; no anterior excavation or lateral lobes; caudally 
directed lateral arms of vinculum strongly sclerotized 
and therefore distinct (in contrast to many other 
nepticulids). Aedeagus a little longer than vinculum 
plate, 195-230 pm long, with small pointed lateral 
extension at the caudal end and with a few (4-5) 
compactly arranged cornuti; cornuti resemble the 
'usual' spine-like ones common in many nepticulids; 
however, in this species they are weakly defined, some- 
times with unsclerotized apices. 

Genitalia ? (Figs 89, 90). Total length about 305- 
365 pm. Abdominal tip widely rounded, anal papillae 
indistinct. Apophyses anteriores distally with longer 
and narrow lateral process and large blunt inner lobe. 
Apophyses posteriores slender and much exceeding 
apophyses anteriores. Ductus spermathecae originat- 
ing from a heavily sclerotized and wrinkled area 
between apophyses posteriores, narrow and long, with 
three sclerotized convolutions. Corpus bursae weakly 
folded, relatively very small, distinctly globular, with 
an oval signum formed by a group of numerous, large, 
blunt pectinations. 

Biology (Figs 100, 101). Host-plant: Psidium 
guajava L. (Myrtaceae). Larvae green, found in 
January; mining leaves. Egg on underside of leaf. Mine 
slender, sinuous, from short to relatively long, mainly 
on upperside of leaf. In the first half of the mine 
blackish frass can fill most of the width, with narrow 
but still distinct clear margins remaining; in the second 
half of the mine frass is deposited as a black narrow 
central line with broad clear margins remaining; some- 
times the central line of frass can be interrupted, but in 
most examples it forms an uninterrupted line. Cocoon 
greyish ochreous, flattened, strongly narrowed to the 
posterior end, ca. 2.5 by 1.5 mm. Voltinism of the 
species is not known. Mining larvae in January were 
found together with old and very old mines from 
probably November-December. Adults from larvae 
collected in January emerged after 20-25 days (in 
February). The new species seems quite common on 
cultivated guava in Ecuador. 

DIAGNOSIS. Very distinctive species; from all 
currently known species of this genus (and most species 
of other genera) it differs strongly by the presence of 
two distinctive forewing fasciae. In the male genitalia 
this species is easily distinguishable by the shape of the 
gnathos (an inverted V). In the female genitalia it 
differs from other Enteucha by the oval signum formed 
by a group of numerous, large, blunt pectinations. The 
unusual host-plant may also make this species distinc- 
tive, but as very little is known about the biology of 
other Enteucha it is uncertain how atypical the utilisa- 
tion of Myrtaceae might be. 

Distribution. Western foothills of the Andes 
(Ecuador), but probably with a much wider distribution. 
Common in the Bucay area (80 km E of Guayaquil) 
(Fig. 101). 

Condition of type material. A few paratype 
specimens from the type series containing 8 specimens 
of this minute species are badly pinned. 

Material examined. Holotype d\ Ecuador: 80 
km E of Guayaquil, Bucay, western foothills of Andes, 
700 m, premontane tropical forest and orchards, 16- 
19.L2001 (Puplesis & Hill), genitalia slide AD0340 



( VPU ). Paratypes, 5 cf , data as holotype, genitalia slides 
AD0341, AD0342, (VPU)], 2 9, genitalia slides 
AD0343, AD0344, forewing venation slide AD0346 
(VPU)]. Mines no. 4722-VPU. 


Stigmella montanotropica Puplesis & Diskus 
sp. n. 

(Figs. 5,29, 30,91,98,99, 102) 

Male. See Remarks. 

Female (Fig. 5). Forewing length: 2.2-2.4 mm. 
Wingspan: 5.0-5. 1 mm. Head: palpi cream: frontal tuft 
from pale orange to bright orange; collar forming two 
clearly separated tufts of cream coloured lamellar 
scales; eyecaps cream; antenna fuscous, comprised of 
22 segments. Thorax, tegulae and forewings fuscous 
brown with some bronzy-golden and purple reflection. 
Cilia dark grey to fuscous. Underside of forewing 
fuscous with some indistinct bronze-golden and pur- 
ple reflection. Hindwing lanceolate, dark brown; cilia 
concolorous. Legs dark fuscous grey, distally cream. 
Abdomen fuscous, shiny, with long anal tufts. 

Genitalia d (Figs 29, 30). Capsule 250-260 pm 
long. Uncus strongly sclerotized, constricted towards 
distal end, with four slender and distinct distal papillae. 
Tegumen very short, simple, band-like. Gnathos with 
large medially narrowed central plate and two very 
slender and long posterior processes parallel with each 
other; anterior extensions of gnathos triangularly lobate, 
weakly developed; lateral arms of gnathos narrowing 
caudally, relatively very short. Valva 144-151 |im 
long, constricted at apex and with two closely spaced 
and pointed apical processes. Transtilla with inwardly 
curved transverse bar, broad triangular angles but with- 
out sublateral processes. Ventral plate of vinculum 
relatively short, one-half or more of length of valva; 
anterior excavation wide but shallow, lateral lobes 
small, distally pointed. Aedeagus broad (90-100 urn) 
and relatively short (21 2-2 1 8 [im long). Vesica with 1 4 
large or very large horn-like cornuti, the largest apical. 
Juxta apparently absent. 

GENITALIA ? (Fig. 9 1 ). Total length about 1 1 25 |im. 
Anal papillae weakly developed, two widely rounded 
but very short setose lobes. Apophyses anteriores 
gradually tapering distally, straight. Apophyses 
posteriores very slender and long, their distal ends 
slightly exceeding apophyses anteriores. Accessory 
sac globular, wrinkled, with slender, medium to weakly 
sclerotized ductus spermathecae, its inner duct with 
about nine neatly arranged convolutions. Corpus bursae 
large, oval, with a semicircular signum comprising 
very numerous pectinations; proximal part of corpus 
bursae long and strongly folded. 

BIOLOGY (Figs 98, 99, 102). Host-plant: Acalypha 
sp. (Euphorbiaceae). Larvae in January-February, 
mining leaves. Egg on upperside or underside of leaf, 
usually deposited close to a small leaf- vein. Mine very 
long, slender, sinuous to contorted, mainly on upperside 
of leaf, more than 100 mm in straightened length. In 
the first half of the mine dark brown or blackish frass 
may fill up the entire width of the mine, latterly depos- 
ited in arcuate waves with or without leaving narrow, 
clear margins to the mine; the longest, final stage of the 
mine is usually with a broad trace of black granulated 
frass, leaving clear margins. Cocoon whitish, oval, 
flattened, strongly narrowed to the posterior end, ca. 
2.5 by 1 .5 mm. Voltinism not known. Mining larvae in 
early February were found together with old and very 
old mines from probably November-January. Currently 
adults are known from February-early March. 

Diagnosis. Belongs to the S. salicis-group (see 
Puplesis, Diskus, Robinson & Onore, 2002). Easily 
recognisable among all other species of the group by 
the combination of smoothly scaled fuscous forewings 
(without fascia) with numerous large cornuti in the 
aedeagus. Among Neotropical representatives of the 
group, the male genitalia most closely resemble those 
of andina (Meyrick) and cuprata (Meyrick); however, 
this species may be distinguished by the uniformly 
dark forewing and hindwing, and by the shape of the 
cornuti and the apical processes of the valva. 

DISTRIBUTION. Tropical montane forest of the west- 
ern slopes of the Andes (Ecuador) (Fig. 102); known 
from altitudes around 1200 m. See also Remarks. 

Condition of type material. Holotype and 
especially female paratype in relatively good condition; 
the male paratype comprises only a genitalia slide 
(29618 BMNH), the genitalia, fully formed and 
sclerotized, dissected from a dead pupa. 

Material examined. Holotype ?, Ecuador: E of 
Santo Domingo de los Colorados, Tandapi, montane 
tropical forest, 1200 m, larvae on leaves of Acalypha 
sp. (Euphorbiaceae), 3-6. ii. 2000 (Puplesis), genitalia 
slide no. 29619 (BMNH). Paratypes, data as holotype, 
Id genitalia slide [only - see above] no. 29618 
(BMNH), 1 9, slide AD0336 (VPU). Mines no. 4628 
(VPU). See also Remarks. 

REMARKS. The description of the external features 
of this species is based on females. 

A few specimens closely resembling this species 
were reared from Acalypha leaf-mines in 2001 in 
premontane tropical forest SE of Tena (Misahualli) 
and montane forest in Banos. These specimens will not 
be dealt with further until additional material is avail- 
able; they may represent a closely related but different 
allopatric new species. The specimens are deposited in 
VPU (male genitalia slides AD 0345, AD 0347, AD 



Stigmella nubimontana Puplesis & Diskus 
sp. n. 

(Figs 6, 31-36, 94,95) 

Male (Fig. 6). Forewing length: about 2.5 mm. Wing- 
span: 5.2-5.3 mm. Head: palpi grey-cream; frontal tuft 
pale orange; collar small, dark brown with some purple 
reflections; eyecaps cream with strong golden reflec- 
tion; antenna fuscous, comprising of 29-30 segments. 
Thorax and tegulae dark brown with golden reflection. 
Forewing coppery-brown with golden and purple 
reflection, purple reflection particularly strong along 
costal margin towards fascia; fascia postmedial, weakly 
demarcated, comprising coppery-golden shining 
scales; area behind fascia dark brown with purple 
iridescence; additionally a slender terminal spot or 
fascia comprising the same shining, coppery-golden 
scales as the fascia. Cilia fuscous to grey at distal end. 
Underside of forewing brown with some purple 
reflection distally. Hindwing and its cilia brown-grey. 
No androconia on hindwing or forewing. Legs fuscous, 
shiny. Abdomen fuscous. 

Female. Unknown. 

Genitalia d (Figs 31-36). Capsule 290-309 pm 
long. Uncus with shallow distal excavation and almost 
parallel, strongly sclerotized lateral margins; the struc- 
ture may vary a little in length. Tegumen small, simple, 
band-like. Gnathos with very long, slender, parallel 
caudal processes, narrowed lateral arms and short, 
wide central plate, gradually constricted at the middle. 
Valva 168-207 pm long, broad, with two inwardly 
curved and pointed apical processes; the ventral proc- 
ess blunt, lobe-like. Transtilla with slender transverse 
bar and almost rounded corners. Juxta apparently 
absent. Ventral plate of vinculum with large triangular 
lobes and deep semicircular excavation distally. 
Aedeagus 195-220 pm long, with many fine spine- 
like cornuti and three large horn-like ones. 

Biology (Figs 94, 95). Host-plant: Rubus sp. 
(Rosaceae). Larvae are known mining leaves in Janu- 
ary-February but the mining period is likely to be 
much longer. Egg on the underside of leaf, usually 
deposited close to a leaf-vein. Mine usually between 
two leaf-veins, a compactly deposited gallery which 
resembles an elongated blotch, about 2.8-3.8 cm in 
straightened length. Beginning of mine slender and 
filled with dark brown frass; later the gallery widening 
considerably, the frass dark brown, deposited in a 
broad compact central line with clear margins; final 
length of mine very broad (sometimes a blotch), with 
granulated dark brown frass deposited more or less 
irregularly or into a relatively slender central line. 
Final part of mine usually covering beginning of mine, 
so the initially slender gallery may be impossible to 
see. Larva yellowish. Cocoon pale brown with some 

yellowish tint, oval, flattened, gradually narrowed to 
the posterior end, ca. 2.8-3.0 x 1.8 mm. Voltinism not 
known. Numerous mining larvae in early February 
were found together with very numerous old and very 
old mines from probably November-January period. 
Adults known from February-early March. 

Diagnosis. Belongs to the S. salicis-gwup (see 
Puplesis, Diskus, Robinson & Onore, 2002); two shin- 
ing forewing fasciae and the combination of three large 
cornuti with numerous fine, spine-like cornuti make 
this a distinctive species of the group. This high Andes 
species is somewhat similar and related to rubeta, the 
other Rubus-mxnmg species from montane tropical 
forest on the western slopes of the Andes; nubimontana 
differs from rubeta in the more strongly developed 
fasciae of the forewing, particularly in the unelaborated 
uncus & tegumen, and the presence of many fine, 
spine-like cornuti (in rubeta no tiny cornuti are present). 
There are some differences in the mines too: the slen- 
der beginning of the gallery made by rubeta has a very 
slender, broken line of blackish frass whereas in mines 
of nubimontana dark brown frass fills up the entire 
width of the gallery. 

Distribution. Cloud forest of the high Andes 
(Ecuador), at about 3500 m, where it appears to be very 

Condition of type material. The male holotype 
is a reared specimen with well-preserved forewings 
and antennae but with detached hindwings stored in a 
small gelatine capsule beneath the specimen. The two 
paratypes came from dead late-stage pupae and are 
therefore not represented by pinned adults, just by 
good-quality genitalia preparations, mines and 

Material examined. Holotype d , Ecuador: Napo 
Region, Andes, 3500 m, Papallacta, cloud forest, larvae 
2-3. ii. 2000 on Rubus sp. (Puplesis & Hill), genitalia 
slide no. 29614 (BMNH). Paratypes, 2c?, data as 
holotype, genitalia slide nos AD032 1 , AD0322 ( VPU). 
Mines no. 4630 (VPU). 

Remarks. During fieldwork in Ecuador in 2000 very 
many mining larvae of this species were collected in 
the Andes (Papallacta). However, for some unknown 
reason (not parasitism - very few parasitoids were 
observed), very few adults emerged. Another species 
mining on the same Rubus sp. at the same time in the 
same locality was also noticed, and a single female 
specimen was reared (no. AD0337 VPU). This last 
species is not included here because of its inadequate 

Stigmella rubeta Puplesis & Diskus sp. n. 

(Figs 7, 37-39, 92, 96, 97, 102) 

MALE (Fig. 7). Forewing length: about 2.0 mm. Wing- 



span: 4.5 mm. Head: palpi grey-cream or brown-cream; 
frontal tuft orange; collar small, fuscous; eyecaps cream 
with some golden reflection; antenna fuscous, number 
of segments unknown. Thorax and tegulae fuscous 
with some golden reflection. Forewing fuscous with 
golden and purple reflections, and without distinct 
fascia: there are only some shiny silver scales in the 
usual area of a median-postmedial fascia and at apex of 
forewing. Cilia fuscous to grey at the distal end. Un- 
derside of forewing brown. Hindwing and its cilia dark 
grey. No androconia on hindwing or forewing. Legs 
fuscous, distally grey-cream. Abdomen fuscous. 

Female. Externally probably very similar to male, 
however, the single female known came from a dead 
pupa and therefore the scaling pattern could not be 

Genitalia d (Figs 37-39). Capsule 290-300 urn 
long. Uncus elaborate in comparison with related 
species: the bilobate, deeply divided structure is paral- 
leled by an additional broad dorsal lobe (Fig. 38). 
Tegumen small, also unusually elaborate, with strongly 
sclerotized anterior extension distally. Gnathos with 
very long, slender, parallel caudal processes, narrowed 
lateral arms and short, wide central plate constricted at 
middle and with a small anterior papilla. Valva 175- 
1 85 urn long, broad, with rounded inner lobe of valva 
(not forming a process as in nubimontana) and with 
inwardly curved and pointed apical process. Transtilla 
with slender transverse bar and angular corners. Juxta 
apparently absent. Ventral plate of vinculum with large 
triangular lobes and deep semicircular excavation 
distally. Aedeagus about 200 urn long, without small, 
spine-like cornuti, but with seven large ones: three of 
them broad-based and four well-sclerotized and slender. 

Genitalia ? (Fig. 92). Total length about 800- 
820 um. Abdominal tip widely rounded. Apophyses 
posteriores about 187-225 um long, very slender and 
straight. Apophyses anteriores slightly longer and 
broader. Accessory sac globular, wrinkled. Ductus 
spermathecae membranous, a short section only coiled 
and strongly sclerotized. Corpus bursae distinctly oval, 
with numerous pectinations all over the sac; proximal 
part of corpus bursae narrowed, short and strongly 

Biology (Figs. 96, 97, 102). Host-plant: Rubus sp. 
(Rosaceae). Larvae mine leaves in January-February 
(but the mining period is probably much longer). Egg 
on underside of leaf, usually deposited close to a leaf- 
vein. Mine usually between two leaf-veins, starting as 
a long and very narrow gallery with blackish or black 
frass deposited in a broken, slender to very slender 
central line; later the gallery widening, sometimes 
very contorted and resembling more a blotch; the 
blackish (sometimes brown) frass in the wider part of 
the mine tending to be deposited in a broad, irregular 

central line. Total length of straightened gallery about 
A.2-A.6 mm (about half of this length belongs to the 
very slender part of the mine), but the mine does not 
superficially appear as long, as the slender region is 
very indistinct; wider part very convoluted, sometimes 
covering beginning of mine. Larva yellowish. Cocoon 
pale brownish with some yellowish tints, oval, flat- 
tened, gradually narrowed to the posterior end, 2.5-2.6 
x 1.8 mm. Voltinism not known. Numerous mining 
larvae were found in early February together with very 
numerous old and very old mines from probably the 
November-January period. Adults are known from 
February-early March. 

Diagnosis. Belongs to the S. salicis-gcoup (see 
Puplesis, Diskus, Robinson & Onore, 2002); this tropi- 
cal montane /?«/;>«.y-mining species is most similar and 
related to nubimontana, the high-Andean species also 
feeding on Rubus. Differs from nubimontana by the 
darker wings, less prominent forewing pattern, the 
elaborate uncus and tegumen, the absence of very 
small spine-like cornuti (which are very characteristic 
for nubimontana) and by the presence of broad-based 
cornuti on the vesica (which are apparently absent in 
nubimontana). Additionally, there are some differ- 
ences in the mines: the slender start of the gallery made 
by rubeta has a very slender, broken line of blackish 
frass (Fig. 97) whereas in mines of nubimontana dark 
brown frass tills the whole width of the gallery (Fig. 

Distribution. Tropical montane forest of the west- 
ern slopes of the Andes (Ecuador), at about 1200 m, 
where it appears to be quite abundant. 

Material examined. Holotype d , Ecuador: west- 
ern Andes, E of Santo Domingo de los Colorados, 
Tandapi, montane tropical forest, larvae 3-6.ii.2000 
on Rubus sp. (Puplesis & Hill), genitalia slide no. 
296 1 6 (BMNH ). Paratype: ? , data as holotype, genita- 
lia slide no. 29617 (BMNH) (Puplesis) (genitalia only 
- see Remarks). Mines no. 4629 (VPU). 

Remarks. The female specimen mentioned above 
was taken from a fully-developed pupa, and the 
paratype is represented only by a preparation of the 
fully formed and sclerotized genitalia. It appears that 
rubeta and nubimontana represent two very closely 
related, allopatric species. 

Stigmella austroamericana Puplesis & Diskus 
sp. n. 

(Figs 8, 40-43) 

MALE (Fig. 8). Forewing length: 1.6-1. 7 mm. Wing- 
span: 3.6-3.8 mm. Head: palpi (and face) whitish 
cream: frontal tuft pale yellowish orange, collar large, 
whitish cream; eyecaps whitish cream; antenna of 
contrasting colour, fuscous on upperside, but distinctly 



cream on underside, comprising of 38 segments. 
Thorax, tegulae and forewing dark grey-brown irrorated 
uniformly with grey with strong golden reflection; a 
few scales just before forewing cilia may appear glossy 
white when viewed at a certain angle. Cilia grey. 
Underside of forewing brownish fuscous: fuscous- 
tipped scales densely irrorated grey-cream on grey 
background. Hindwing and cilia brown-grey. No 
androconia on forewing or hindwing. Legs brownish 
cream distally and on underside, elsewhere dark grey- 
brown. Abdomen silver-blackish on upperside, grey, 
glossy; tufts indistinct. 

Female. Unknown. 

Genitalia <S (Figs 40-43). Capsule about 320 um 
long. Uncus comprising two very large, distally rounded 
lobes. Tegumen band-shaped, slightly spined distally. 
Gnathos with two very slender and long caudal 
processes and pair of anterior processes that are 3x 
broader and about one-half length of anterior processes; 
central plate very slender; lateral arms short and narrow. 
Valva 190-196 urn long, bulged medially and 
constricted to a pointed apical process caudally, with 
long plumose scales in apical half. Transtilla with 
sublaterally directed processes and transverse bar which 
is not fused in the middle. Ventral plate of vinculum 
with deep, semicircular anterior excavation and long, 
distally truncate lateral lobes. Aedeagus 215-226 urn 
long, slender, with tiny, pointed apical processes on 
both sides of the tube, and with a few fine, spine-like 
cornuti in the apical part. Juxta membranous, lobate, 
hardly visible. 

Biology. Adults collected in late January. 

DIAGNOSIS. Belongs to the same species-group (see 
Remarks) as two other Neotropical species: 
plumosetaeella Newton & Wilkinson and barbata 
Puplesis & Robinson. Externally this species is easily 
distinguished from the fasciated plumosetaeella by the 
dark forewing; it is externally similar to barbata but 
differs in the absence of brownish androconial scales at 
the apex of forewing and in the absence of long thick- 
ened brown androconial scales on the dorsal margin of 
the hindwing. In the male genitalia austroamericana is 
easily distinguished from both related species by the 
long lobes of the vinculum, the huge, deeply lobate 
uncus, and long, almost straight apical processes of the 

Distribution. Ecuador; Amazon premontane rain- 
forest, at altitudes of about 500 m. 

Condition of type material. The holotype is well 

Material examined. Holotype d , Ecuador: Napo 
Region, SE of Tena, near Rio Napo, Jatun Sacha, 
Amazon rainforest, 500 m, 26-3 l.i. 2000 (Puplesis & 
Hill), genitalia slide no. 29613 (BMNH). 

Remarks. This species represents a very distinctive 
member of a still unnamed species-group obviously 
comprising also plumosetaeella Newton & Wilkinson 
and barbata Puplesis & Robinson; on some similari- 
ties in forewing pattern and genitalia it is also easy to 
assume that austroamericana is more related to barbata 
Puplesis & Robinson; they may represent allopatric 


Fomoria repanda Puplesis & Diskus sp. n. 

(Figs 9, 44-52) 

MALE (Fig. 9). Forewing length: 1.6- 1.9 mm. Wing- 
span: 3.7^4.4 mm. Head: palpi cream to ochreous 
cream; frontal tuft ochreous orange; collar indistinct, 
comprising pale ochreous piliform scales; eyecaps 
cream with or without some pale brownish scales; 
antenna brownish to brown or ochreous brown on 
upperside, cream on underside, comprising of 24-28 
segments. Thorax and tegulae ochreous cream with 
many brownish, brown or dark brown scales; these 
brown scales may be weakly developed and/or easily 
lost. Cream to brownish cream background of forewing 
irrorated with brown and dark brown scales; these dark 
scales more densely distributed on base of wing, close 
to costa and especially distinctive and dark behind the 
fascia, i.e., in apical part of forewing. Fascia median- 
postmedial, broad and oblique, formed by 
fuscous-brown scales. Cilia ochreous cream. Under- 
side of forewing yellowish brown. Hindwing and cilia 
cream. No androconia on hindwing or forewing. Legs 
cream with or without some dark brown shading. 
Abdomen yellowish brown on upperside, paler on 

Female. Unknown. 

Genitalia d (Figs 44-52). Capsule 330-342 urn 
long. Pseuduncus distinctly bilobed; the lobes very 
large, irregularly triangular. Uncus with distally 
rounded median lobe (see Remarks). Gnathos with 
relatively short caudal element and long slender lateral 
arms. Valva 100-109 um long, divided into slender, 
distally rounded and setose lobe, and slender and 
distally pointed dorsal lobe. Transtilla with very 
extended lateral arms and with an angular plate (trans- 
verse bar); plate of transtilla weakly sclerotized. 
Vinculum very long, without anterior (distal) excava- 
tion or lobes. Juxta apparently absent. Aedeagus very 
large, about 575 um long, with a wide band of broad, 
triangular and lobe-like cornuti, and with about seven 
very large horn-like cornuti in apical one-quarter; three 
orfour of these horn-like cornuti straight, remainder 
with curved bases. Aedeagus of holotype (Fig. 45) 
with inverted vesica, and not all cornuti preserved and 
visible; see paratype (Fig. 52). 



Biology. Adults fly in January. 

DIAGNOSIS. This distinctive Amazon rainforest 
species forms a natural group with a few other 
Neotropical Fomoria: molybditis (Zeller) [Colombia], 
diskusi Puplesis & Robinson [Belize], and unnamed 
species 29122 [Belize] (see Puplesis & Robinson, 
2000). Externally repanda differs from all these species 
in the dark fascia of the forewing. The male genitalia 
exhibit some similarities to molybditis and species 
29122 in the presence of numerous cornuti in the 
aedeagus. However, repanda clearly differs from 
molybditis in the deeply divided valva without a spine- 
like process and in the broad median lobe (element) of 
the uncus; from species 29122 it differs in the very 
long aedeagus, slender ventral lobe of the valva, and 
broad uncus. 


Lowland Amazon rainforest (Ecua- 

paratype well preserved. 

Holotype and 

Material examined. Holotype d , Ecuador: Napo 
Region, SE of Coca, near Rio Tiputini, Yasuni National 
Park, 260 m, 15-25.i.2000 (Puplesis & Hill), genitalia 
slide no. 29603 (BMNH). Paratype: d\data as holotype, 
genitalia slide no. AD0315 (VPU), 

REMARKS. In the genitalia slides of both currently 
known specimens the median lobe of the uncus is 
directed anteriorly, but it is uncertain that this is the 
natural position and not an artefact of preparation. The 
normal position might well be with the median lobe 
directed caudally, as is characteristic of other Fomoria. 

Fomoria tabulosa Puplesis & Diskus sp. n. 

(Figs 10.53-55) 

Male (Fig. 10). Forewing length: 2.6 mm. Wing- 
span: about 5.6 mm. Head: palpi cream with some 
brownish shading; frontal tuft pale ochreous; collar 
inconspicuous, comprised of ochreous cream pilifonn 
scales; eyecaps large, yellowish cream; antenna ochre- 
ous cream distinctly annulated with brown, comprising 
of about 35 segments. Thorax, tegulae and forewing 
uniform: greyish cream background densely irrorated 
with brown and dark brown scales. Cilia and underside 
of forewing brownish. Hindwing lanceolate, brown- 
ish; its cilia also brownish. No androconia on forewing 
or hindwing. Legs pale ochreous distally with fuscous 
shading. Abdomen brown; tufts weakly developed. 

Female. Unknown. 

Genitalia d (Figs 53-55). Capsule 408-415 u.m 
long. Pseuduncus triangular with a short, slender and 
distally truncate process. Uncus inverted 'V'-shaped, 
with short and blunt extension, which in most Fomoria 

is caudally directed [the extension of the uncus in a 
ventral direction in the holotype - see Fig. 53 - was 
probably caused by the process of fixation and 
mounting in Euparal]. Gnathos with single, slender 
caudal process and slender, caudally directed lateral 
arms. Valva 240-253 |im long, broad at base, strongly 
narrowed towards apex, very slender and slightly curved 
inwardly in apical third. Transtilla with slender trans- 
verse bar and short, slender sublateral processes. Juxta 
(Fig. 54) represented by a pair of lateral, lobe-like, 
spinose processes behind (dorsal to) ventral lobes of 
valvae. Ventral plate of vinculum half length of valva, 
narrowed distally and with small rounded sublateral 
corners; distal excavation of vinculum plate particu- 
larly shallow. Aedeagus 332 um long, with broad 
inverted 'U'-shaped sclerite; no apical carinae or spine- 
like cornuti. 

Biology. Adults collected in January. 

DIAGNOSIS. Easily distinguished from other Fomoria 
and all other nepticulids by the inverted 'U'-shaped 
sclerite in the aedeagus. 

Distribution. Lowland Amazon rainforest (Ecua- 

Condition of type material. The holotype is in 
good condition. 

Material examined. Holotype d, Ecuador: Napo 
Region, SE of Coca, near Rio Tiputini. Yasuni National 
Park, 260 m, 1 5-25.i.2000 (Puplesis & Hill), genitalia 
slide no. 29625 (BMNH). 


Acalyptris ecuadoriana Puplesis & Diskus 
sp. n. 

(Figs 11,56-58) 

MALE (Fig. 11). Forewing length: 2.5 mm. Wing- 
span: 5.4-5.5 mm. Head: palpi cream; frontal tuft 
ochreous; collar indistinct, consisting of ochreous 
cream piliform scales; eyecaps large, cream, with some 
brownish scales; antenna on upperside broadly annu- 
lated with brown to almost cream, cream on underside, 
comprising of about 28 segments. Thorax, tegulae and 
forewing before the fascia golden cream with some 
brownish scales close to forewing costa. Fascia post- 
medial, oblique, formed by fuscous brown scales; area 
behind fascia brown-grey with some fuscous-brown 
tipped scales. Cilia cream. Some long, thickened, dark 
brown scales extending over hindwing from dorsal 
margin of forewing. Underside of forewing brownish 
cream except margins which are brown. Hindwing 
lanceolate, neatly covered with small brownish (prob- 
ably androconial) scales (on upperside only); cilia of 
hindwing brownish or greyish. Legs ochreous cream. 



Abdomen dark brown to yellowish brown with some 
dark brown scales; on upperside - brownish to yellow- 

Female. Unknown. 

Genitalia d (Figs 56-58). Capsule 280 urn long. 
Pseuduncus triangular and neatly papillated distally. 
Uncus paired, X-shaped, broadened anteriorly. Gnathos 
(Fig. 57) with slender, bar-like central plate, and very 
elaborate, unusual lateral arms, their ventrally 
sclerotized margins resembling distinctly pointed 
caudal processes; ventral arms of gnathos with inwardly 
directed dorsal lobes; real caudal process(es) absent. 
Tegumen not forming lateral rod-like sclerites beneath 
valvae, along capsule as in many other Neotropical 
Acalyptris. Valva 168-173 urn long, broad in basal 
two-thirds and very slender, inwardly curved in apical 
third. Transtilla with narrow, medially very constricted 
(or not fused) transverse bar and outwardly curved 
sublateral processes one-half length of transtilla. Juxta 
a sclerotized transverse bar behind valval bases, slightly 
varying in shape. Ventral plate of vinculum with widely 
rounded lateral lobes and very shallow anterior exca- 
vation. Aedeagus 244 urn long, sclerotized and 
distinctly constricted at base, and with three very long 
horn-like sclerites in apical half; one of them with a 
strongly sclerotized basal extension. 

Biology. Adults collected in January. 

Diagnosis. Together with the Amazonian onorei 
and the Central American latipennata Puplesis & 
Robinson and dividua Puplesis & Robinson this species 
forms a distinctive species group. Externally 
ecuadoriana clearly differs from onorei in the well- 
developed forewing fascia, the absence of androconial 
patches on the hindwing and by the lanceolate hindwing 
shape; in onorei the hindwings are distinctly broad- 
ened. From latipennata and dividua it differs in the 
presence of long piliform scales on the dorsal margin 
of the forewing and by the background colour of the 
forewing - latipennata is paler, and dividua darker. In 
the male genitalia this species differs from all related 
ones in the absence of inner, spine-like valval proc- 
esses, a distinctly X-shaped uncus with broadened 
anterior components, distally excavated vinculum, very 
broad valva in basal two-thirds, and the specialized 
shape of the gnathos. 


Premontane Amazon rainforest 

Condition of type material. The type series is 

Material examined. Holotype d , Ecuador: Napo 
Region, near Rio Napo, Jatun Sacha, premontane rain- 
forest, 400-500 m, 26-31.i.2000 (Puplesis & Hill), 
genitalia slide no. 29624 (BMNH). Paratypes: Id, 

data as holotype, genitalia slide no. AD0325 (VPU); 
Id, data as holotype but Misahualli, 22-31.i.2001 
(Puplesis & Hill), genitalia slide no. AD0349 (VPU). 

Acalyptris onorei Puplesis & Diskus sp. n. 

(Figsl, 12, 13,59-64) 

Male (Figs 1, 12, 13). Forewing length: 2.0-2.2 
mm. Wingspan: 4.5-4.9 mm. Head: palpi cream; fron- 
tal tuft ochreous orange; collar indistinct, comprised of 
ochreous cream piliform scales; eyecaps cream; 
antenna cream, annulated with brownish to fuscous 
brown on upperside, comprising of 26-28 segments. 
Thorax and tegulae irrorated with numerous brownish 
to brown scales. Forewing broad with some bluish and 
green reflection; in basal three-fifths, cream to grey- 
cream background indistinctly, more or less smoothly 
irrorated with brownish scales, apical two-fifths 
distinctly irrorated with larger, brown to fuscous brown 
scales; a clear fascia undeveloped, however, numerous 
fuscous brown scales distributed just behind middle of 
forewing resemble an irregular, oblique postmedial 
fascia. Cilia greyish cream. Underside of forewing 
dark brown with very long, median band of cream 
androconial scales along width of wing. Forewing 
venation (Fig. 1 ) typical for Acalyptris, however R4 
and R5 preserved as two separate veins in contrast to 
the usual Acalyptris venation where these veins are 
fused. Hindwing (Fig. 13) unusually broad, on 
upperside with large, elongated patch of neatly arranged 
cream scales close to hind margin and base; covered 
with brown scales, very densely deposited just before 
apex; apex brownish; with single frenulum and, close 
to base, a dark brown hair pencil comprising many 
very long, thickened chaetae-like scales; behind hair 
pencil with many cream, elongated lamellar scales 
arising from anterior margin of wing. Cilia of hindwing 
cream, overlapped by long, thickened, brownish scales 
from posterior margin near apex. Underside of 
hindwing brownish to brown; sometimes with a paler 
zone along main vein. Legs pale ochreous or ochreous 
cream. Abdomen brown with broad dark bands along 
upperside, underside yellowish brown or brownish. 

Female. Unknown. 

Genitalia d (Figs 59-64). Capsule 295 urn long. 
Pseuduncus with two small triangular lobes apically; 
excavation between apical lobes as deep/broad as the 
length/width of each lobe. Uncus (Fig. 62) paired, 
comprising lateral, strongly sclerotized plates which 
are widened at distal end and outwardly curved at 
bases. Gnathos (Fig. 63) paired, comprising two lateral, 
strongly sclerotized, broad, V-shaped sclerites; no 
central plate or any transverse bar connecting lateral 
parts of gnathos. Tegumen not forming lateral rod-like 
sclerites beneath valvae, along capsule as in many 
other Neotropical Acalyptris. Valva 1 75-1 8 1 urn long. 



with large, horn-like apical process and similar medial 
one. A pair of small rounded sclerotizations visible on 
diaphragma between medial process of valva and 
transtilla bar; homology of these sclerites unknown. 
Transtilla with narrow, long, bridge-like transverse 
bar, and outwardly curved sublateral processes one- 
third length of bar. Juxta apparently absent. Vinculum 
large, ventral plate trapezoidal, gradually narrowed 
towards anterior (distal) end, without lateral lobes and 
without anterior excavation. Aedeagus about 255 u.m 
long, strongly narrowed in basal third, and with three 
large, horn-like sclerites apically. 

Biology. Adults collected in January. 

Diagnosis. Together with the Amazonian 
ecuadoriana and two Central American species, 
latipennata Puplesis & Robinson and dividua Puplesis 
& Robinson, onorei forms a distinctive species group. 
Externally it differs from these related species in the 
distinctly broadened hindwing with androconial 
patches, and weakly developed (or undeveloped) 
forewing fascia. In the male genitalia onorei differs 
from all related species by the unique gnathos 
comprised of two lateral sclerites without a transverse 


Lowland Amazon rainforest 

Condition of type material. The holoty pe is well 
preserved; the paratype has a slightly rubbed head and 

Material examined. Holotype <S . Ecuador: Napo 
Region, SE of Coca, near Rio Tiputini, Yasuni National 
Park, 260 m, 1 5-25.i.2000 (Puplesis & Hill), genitalia 
slide no. 29623 (BMNH). Paratype <$: data as holotype, 
genitalia slide no. AD0324 (VPU), forewing venation 
slide no. AD0351 (VPU). 

Remarks. This remarkable species is named in 
honour of Dr. Giovanni Onore (Departamento de 
Biologia, PontificiaUniversidadCatolica del Ecuador, 
Quito, Ecuador) who helped to organize the first 
collecting trip to Ecuador specifically targeted at Ama- 
zon rainforest Nepticuloidea. 

Another species, closely related to onorei, was found 
by Puplesis & Hill (2001) in premontane tropical 
rainforest (Misahualli, 17 km SE of Tena). The speci- 
men will not be described until further material becomes 
available; it is deposited at VPU (genitalia slide no. 

Acalyptris basihastatus Puplesis & Diskus 
sp. n. 

(Figs 14, 65-67, 71) 

MALE (Fig. 14). Forewing length 1.9-2.1 mm. Wing- 
span: 4.3-4.6 mm. Head: palpi cream; frontal tuft 

ochreous to pale orange-ochreous; collar indistinct, 
comprised of cream piliform scales; eyecaps cream; 
antenna brownish, comprising of about 33-35 
segments. Thorax and tegulae densely irrorated with 
brown (or dark brown) scales. Golden cream back- 
ground of forewing (except costal area) irrorated with 
small, indistinct pale brownish scales; the narrow area 
along costa densely covered with dark brown scales; 
apical part of forewing irrorated with dark brown or 
fuscous-tipped scales; some dark scales overlapping 
on cilia. Cilia cream. Underside of forewing dark 
brown. Hindwing lanceolate, brownish or greyish; cilia 
greyish. No androconial patches on hindwing or 
forewing. Legs cream with golden reflection. Abdomen 
dark brown on upperside and yellowish brown on 
underside, with paired abdominal tuft of long piliform 

Female. Unknown. 

Genitalia d (Figs 65-67. 71). Capsule 312-320 
u.m long. Tegumen extended into distally bilobed 
pseuduncus; each lobe rounded and papillated. Uncus 
in shape of inverted Y; the caudal process slender and 
short. Gnathos with large caudal process, somewhat 
slender lateral arms, and subangular central plate. 
Tegumen forming lateral rod-like sclerites beneath 
valvae, along capsule. Valva (Fig. 71) 205-218 (im 
long, with large, pointed, subcaudally directed basal 
process, bulged inner lobe and gradually constricted 
apical half. Transtilla with slender and long transverse 
bar and similarly slender sublateral processes one-half 
length of bar. Juxta indistinct or apparently absent. 
Ventral plate of vinculum broad and short, with very 
broad, triangular, distally rounded lateral lobes. 
Aedeagus 439^148 |im long (including distal carina); 
tube equally broad, apically with five large, mostly 
asymmetrical carinae and large, elongated, semi- 
triangular sclerotization near bases of lowest (lateral) 
carinae; in undissected genitalia, this elongated plate- 
like sclerotization probably attaches to caudal margin 
of vinculum. Vesica with some very small triangular 
cornuti close to the elongated, plate-like sclerotization. 
Cathrema very sclerotized, relatively very long, almost 
as long as any carinae. 

Biology. Adults collected in late January. 

Diagnosis. This species is distinguished from all 
currently known Neotropical Acalyptris by the large, 
straight and pointed, distinctly basal process of the 
valva. Externally it might be confused with other 
Acalyptris with irrorated forewings, particularly with 
the central american martinheringi and the related, 
Amazonian pseudohastatus or artiatlosus. In 
martinheringi the somewhat similar basal processes 
are slightly curved or bifid, and are associated with the 
juxta, not the valva; the pseuduncus is distally rounded, 
and the valva slender. In pseudohastatus the valval 



process is attached for its entire length to the inner lobe 
of the val va and is sinuous, not straight; the pseuduncus 
is truncated, not bilobed; the aedeagal carinae are very 
long. In articulosus, in contrast to basihastatus, the 
basal lobe of the valva is articulated. 

Distribution. Amazon premontane rainforest 
(Ecuador), at altitudes of about 500 m. 

Condition of type material. The holotype and 
four paratypes are in good or satisfactory condition, 
with spread wings, and are little worn. The best 
preserved scaling of forewing and head is undoubtedly 
in paratype AD0316. 

Material examined. Holotype d , Ecuador: Napo 
Region, near Rio Napo, Jatun Sacha, premontane rain- 
forest, 400-500 m, 26-3U.2000 (Puplesis & Hill), 
genitalia slide no. 29605 (BMNH). Paratypes: 3d", 
data as holotype, genitalia slide nos 29606 (BMNH), 
AD0316(VPU),AD0317(VPU); Id 1 , data as holotype 
but Misahualli, 22-3 l.i. 2001 (Puplesis & Hill). 
genitalia slide no. AD0350 (VPU). 

Acalyptris pseudohastatus Puplesis & Diskus 
sp. n. 

(Figs 15,68-70,72) 

MALE(Fig. 15). Forewing length: 2. 1-2.2 mm. Wing- 
span: 4.7-4.9 mm. Head: palpi cream; frontal tuft 
ochreous orange to pale ochreous; collar indistinct, 
comprised of cream piliform scales; eyecaps cream; 
antenna brown to brownish, comprising of about 36-38 
segments. Thorax and tegulae densely irrorated with 
brown (or dark brown) scales. Greyish cream or golden 
cream background of forewing irrorated with brownish, 
brown and dark brown scales; dark brown-tipped scales 
particularly abundant in apical half of forewing; some 
dark scales overlapping on forewing cilia; a narrow area 
along costa densely covered with dark brown scales. 
Cilia greyish. Underside of forewing brown to dark 
brown. Hindwing lanceolate, grey or brownish; cilia 
grey. No androconial patches on hindwing or forewing. 
Legs brownish cream with dark brown shading (particu- 
larly on forelegs). Abdomen dark brown on upperside 
and yellowish brown on underside, with paired abdomi- 
nal tuft of long piliform scales. 

FEMALE. Unknown. 

Genitalia d (Figs 68-70, 72). Capsule 340-354 urn 
long. Tegumen extended into trapezoidal, distally trun- 
cate pseuduncus. Uncus in shapeofinvertedY;thecaudal 
process slender and short. Gnathos with large caudal 
process, slightly slender lateral arms, and with small 
central plate. Tegumen forming lateral rod-like sclerites 
beneath valvae, along capsule. Valva (Fig. 72) 240-249 
urn long, with large, pointed, caudally directed median 
process, bulged inner lobe and gradually constricted 

apical half. Transtilla with slender and long transverse 
bar and very slender sublateral processes half length of 
bar. Juxta indistinct or apparently absent. Ventral plate of 
vinculum broad and very short, with triangular, short, 
distally almost pointed lateral lobes. Aedeagus 350-362 
urn long (including distal carina); tube equally broad, 
apically with five very large carinae and some very small 
triangular cornuti on vesica. Cathrema well sclerotized 
and half as long as any apical carinae. 

Biology. Adults collected in late January. 

Diagnosis. This species is very easily distinguished 
from other Neotropical Acalyptris by the combination 
of the very characteristic median process of the valva 
with the truncate pseuduncus and five very long 
aedaegal carinae. Relationships are to other Amazo- 
nian Acalyptris, notably basihastatus; however, in 
pseudohastatus the valval process is attached for its 
entire length to the inner lobe of the valva and is 
sinuous, not straight; the pseuduncus is truncate (not 
bilobed), the carinae of the aedeagus are very long and 
very slender distally. 

Externally it could be easily confused with other 
Acalyptris with a similarly irrorated forewing, particu- 
larly the central american martinheringi and 
Amazonian basihastatus and articulosus. 

DISTRIBUTION. Amazon premontane rainforest 
(Ecuador), at altitudes of about 500 m. 

Condition of type material. The holotype and 
one paratype (no. AD03 1 9 VPU) are in good condition, 
while the two remaining paratypes are more worn (no. 
29608 BMNH and no. AD03 1 6 VPU). 

Material exam ined. Holotype d , Ecuador: Napo 
Region, near Rio Napo, Jatun Sacha, premontane rain- 
forest, 500 m, 26-3 1 .i.2000 (Puplesis & Hill), genitalia 
slide no. 29607 (BMNH). Paratypes, 3d", data as 
holotype, genitalia slides no. 29608 (BMNH), no. 
AD0318 (VPU), no. AD0319 (VPU). 

Acalyptris articulosus Puplesis & Diskus sp. n. 

(Figs 16,73-75) 

MALE(Fig. 16). Forewing length: 1.8-2.0 mm. Wing- 
span: 4.3^.5 mm. Head: palpi cream; frontal tuft dark 
orange or dull orange; collar comprising orange-cream 
piliform scales; eyecaps cream; antenna brownish to 
brownish cream, comprising of about 34-36 segments. 
Thorax and tegulae irrorated with brown (or dark 
brown) scales. Cream background of forewing irregu- 
larly irrorated with indistinct, small, pale brown scales, 
densest in basal half of wing, and with brown to 
fuscous brown scales predominating in apical half of 
forewing; no fascia or distinct spots, but a few dark 
fuscous scales together may form a patch, however, 
this patch not obviously developed on other wing of 



same specimen; some dark scales overlapping on 
forewing cilia. Cilia brownish cream. Yellowish cream 
background of underside of forewing densely covered 
with small elongated dark brown scales. Hindwing 
generally slender, slightly broadened anteriorly in basal 
half, uniformly covered either with grey-cream or 
brownish grey scales; cilia of hindwing greyish. No 
androconial patches on hindwing or forewing. Legs 
brownish cream to cream with or without fuscous 
shading distally. Abdomen brown on upperside and 
yellowish brown on underside, with paired abdominal 
tuft of long piliform scales. 

Female. Unknown. 

Genitalia d (Figs 73-75). Capsule 270-276 urn 
long. Tegumen caudally extended into a gradually 
tapered pseuduncus, with two slender and short distal 
lobes. Uncus inverted 'Y'-shaped, with a short, slen- 
der, apically thickened and slightly broadened caudal 
process. Gnathos with short, strongly sclerotized lat- 
eral arms; central plate relatively large, laterocaudally 
very strongly sclerotized, anteriorly with more or less 
membranous, triangular extension. Val va 2 1 6-229 urn 
long, slender for entire length, with caudally directed 
basal lobe; lobe divided caudally into two unequally 
developed and unequally sclerotized processes. 
Transtilla with long transverse bar and short, very 
slender sublateral process. Without the lateral rod-like 
sclerites beneath valvae and along capsule, character- 
istic of very many Acalyptris, particularly Neotropical 
species. Ventral plate of vinculum very short, gradually 
constricted distally, with two very small, rounded lat- 
eral lobes with very shallow excavation between them. 
Aedeagus 287-31 7 um long, very broad (1 14-126 uni), 
with a pair of curved, horn-like lateral carinae joined 
by a common transverse bar, five or four (minimum 
three) large, horn-like cornuti, and numerous tiny, 
spine-like triangular cornuti; cathrema strongly 
sclerotized, short and very broad. 

BIOLOGY. Adults collected in late January. 

Diagnosis. This species is easily distinguished from 
other Acalyptris by the basal lobe of the valva which is 
clearly and deeply divided into two unequally devel- 
oped processes, and by the combination of a bilobed 
pseuduncus with a broadened central gnathos plate, 
and tiny vinculum. Externally this species could be 
easily confused with some other Acalyptris possessing 
similarly irrorated forewings, particularly with the 
Central American martinheringi, and the Amazonian 
basihastatus and pseudohastatus (see above). 

Distribution. Amazon premontane rainforest 
(Ecuador), at altitudes of about 500 m. 

Condition of type material. The holotype is in 
fair condition, however the pattern of the forewings is 
not very distinct; the specimen with the best preserved 

scaling and the most distinct forewing pattern is 
paratype no. 29609 BMNH; however, the head of this 
specimen is detached and glued to a separate minuten 
pin. The remaining paratype, no. AD0320 VPU is in 
fair condition, but the hindwings are detached and 
stored in a small gelatine capsule beneath the specimen. 

Material examined. Holotype d\ Ecuador: Napo 
Region, near Rio Napo. Jatun Sacha, premontane rain- 
forest, 500 m, 26-3 1 .i.2000 (Puplesis & Hill), genitalia 
slide no. 29610 (BMNH). Paratypes: 2d, data as 
holotype, genitalia slides no. 29609 (BMNH), no. 
AD0320 (VPU). 

Acalyptris rotundus Puplesis & Diskus sp. n. 

(Figs 17,76-78) 

MALE (Fig. 17). Forewing length: about 1.9 mm. 
Wingspan: about 4.3^1.4 mm. Head: palpi cream; 
frontal tuft ochreous orange or orange; collar incon- 
spicuous, comprised of orange-cream piliform scales; 
eyecaps cream; antenna mostly brownish cream or 
cream with some dark brown shading, comprising of 
about 33-34 segments. Thorax and tegulae pale creamy 
brown, indistinctly irrorated with brownish scales. 
Brownish cream background of forewing irrorated 
with brownish and brown scales before middle, and 
with predominantly larger fuscous scales in apical half 
or two-fifths; forewing pattern tending to be irregular, 
but with median, broad, ill-defined fascia formed by 
whitish cream scales; a few whitish scales also form- 
ing an indistinct, small, more or less rounded apical 
spot that is unequally developed on both wings of the 
same specimen. Cilia distinctly grey, except distal 
area. Underside of forewing brown. Hindwing lanceo- 
late, greyish or brownish; its cilia brownish. No 
androconial patches on hindwing or forewing. Legs 
dark cream. Abdomen with a distinct cream abdominal 
tuft of long piliform scales, dark brown on upperside 
and yellowish brown on underside. 

Female. Unknown. 

Genitalia d (Figs 76-78). Capsule 204-224 um 
long, distinctly rounded. Pseuduncus distinctly 
rounded, strongly sclerotized and papillated. Uncus 
broad, inverted 'Y'-shaped with two very strongly 
sclerotized, lateral, slightly anteriorly curved and 
distally pointed processes, almost as long as caudal 
process; caudal process well-sclerotized, slender. 
Gnathos with large caudal process, broad but short 
lateral arms and relatively large central plate excavated 
from anterior margin. Valva about 120-125 um long, 
generally narrow, slightly broadened towards base, 
with many papillae on inner margin. Both valvae joined 
basally by ventral transverse bar which forms a central 
triangular sclerotization. Transtilla with very narrow 
transverse bar and tiny, very slender sublateral proc- 



esses. Well-sclerotized lateral rod-like sclerites be- 
neath valvae, along capsule, as is characteristic of very 
many Acalyptris species, particularly Neotropical ones. 
Ventral plate of vinculum extremely small, band-like, 
distally truncate or rounded, without any lateral lobes 
and distal excavation. Aedeagus about 100 pm long, 
broad and short, with two lateral, long, very slender 
and pointed carinae, and very many small, fine cornuti. 

Biology. Adults collected in late January. 

Diagnosis. This species is very easily distinguished 
from other Acalyptris by the combination of the very 
characteristic, three-processed uncus and the distinctly 
rounded capsule, and the broad, rounded pseuduncus 
and broad aedeagus with two very slender lateral carinae. 
While the unusual cream forewing fascia of this species 
may be thought of as distinctive, the character is of 
limited value since it is not well-defined and even slight 
abrasion may render it very hard to recognise. 

Distribution. Amazon premontane rainforest 
(Ecuador), at altitudes of about 500 m. 

Condition of type material. The holotype is in 
quite good condition, with well-preserved scaling; the 
wings are not spread but the hindwing is easily visible. 

Material examined. Holotype d , Ecuador: Napo 
Region, near Rio Napo, Jatun Sacha, premontane rain- 
forest, 500 m, 26-3 l.i.2000 (Puplesis & Hill), genitalia 
slide no. 29611 (BMNH). 

Acalyptris amazonius Puplesis & Diskus sp. n. 

(Figs 18, 79-82, 93) 

Male (Fig. 18). Forewing length: 1.5 mm. Wing- 
span: 3.4-3.5 mm. Head: palpi yellowish cream; frontal 
tuft orange-ochreous, distally ochreous brown; collar 
indistinct, comprised of pale ochreous or ochreous 
cream piliform scales; eyecaps yellowish cream; 
antenna dark brown on upperside and yellowish cream 
on underside, comprising of 24-26 segments. Thorax 
and tegulae dark brown, unicolorous. Forewing gener- 
ally dark, basal three-fifths brown, then with an 
indistinct, oblique, fascia-like, paler, brownish-cream 
area; apical area of forewing densely irrorated with 
fuscous brown scales. Cilia pale brown. Underside of 
forewing dark brown or brown. Hindwing and cilia 
grey-brown. No androconia on hindwing or forewing. 
Legs brownish cream with abundant brown to fuscous 
brown shading laterally and distally. Abdomen fuscous 
brown, very reflective on upperside and with brown 
gloss on underside; pair of anal tufts distinct, composed 
of long piliform scales. 

Female (see Remarks). Forewing length: 1.6-1.7 
mm. Wingspan: 3.7 mm. Head: palpi whitish; frontal 
tuft whitish cream; collar sometimes indistinct, com- 
prised of cream piliform scales; eyecaps cream, with 

some brownish scales; antenna brown on upperside 
and brownish on underside, comprising of about 27- 
28 segments. Thorax and tegulae dark brown, 
unicolorous. Forewing generally dark, basal three- 
fifths brown, apically irrorated with fuscous brown 
scales; pale area before the apical region, characteris- 
tic in the male, absent. Cilia brown. Underside of 
forewing brown. Hindwing and cilia grey-brown. Legs 
brownish cream with some brown to fuscous brown 
shading. Abdomen fuscous on upperside, with reddish 
and purple reflection and pale silvery on underside. 

Genitalia 6" (Figs 79-82). Capsule 240-250 urn 
long. Tegumen extended into broad, distally widely- 
rounded and heavily papillated pseuduncus; anteriorly 
with unusual trapezoidal extension. Uncus (Fig. 81) 
with lateral band-like components and caudal element 
directed ventrally; the last very strongly sclerotized 
and resembling a crown. Gnathos with small pointed 
caudal process, small oval central plate and well devel- 
oped lateral lobes wider than the caudal element. Valva 
134-139 pm long, equally narrow for entire length 
except apex which is visibly constricted and very 
strongly papillated; basally valva strongly sclerotized. 
Transtilla with slender transverse bar and sublateral 
processes one-half length of bar. Juxta apparently 
absent. Ventral plate of vinculum half length of valva, 
slightly and gradually constricted distally, and with 
shallow anterior excavation; lateral lobes weakly 
developed, broadly triangular. Lateral rod-like sclerites 
(usually very characteristic for Neotropical Acalyptris) 
distinct and sclerotized only distally while at bases not 
demarcated from tegumen. Aedeagus about 263 pm 
long, with pair of slender lateral carinae, long and 
well-sclerotized cathrema, four slender and long spine- 
like cornuti, and a few very tiny, indistinct ones. 

Genitalia ? (Fig. 93) (see Remarks). Total length 
about 702-708 pm. Abdominal tip broadly rounded. 
Anal papillae undeveloped, but with a distinct, 
sclerotized plate connected with bases of apophyses 
anteriores. Both anterior and posterior apophyses very 
slender, but anterior apophyses one-third to one-quarter 
longer (125-138 pm). Vestibulum with large Y-shaped 
sclerite. Corpus bursae large, elongate-oval with a 
distinct, band-like signum on each side, with tiny 
pectinations around the signa. Accessory sac undevel- 
oped; ductus spermathecae with 8-9 compactly 
arranged convolutions. 

Biology. Adults collected in January. 

Diagnosis. Among Neotropical Acalyptris this 
species resembles externally only the dark Central 
American hispidus Puplesis & Robinson, novenarius 
Puplesis & Robinson and platygnathos Puplesis & 
Robinson; however, it is easily distinguished by the 

In the male genitalia amazonius differs very strongly 



by the crown-shaped uncus and by the apically slightly 
constricted but heavily papillated valva. 

Distribution. Lowland Amazon rainforest 

Condition oftype material. The holotype is well 

Material examined. Holotype d\ Ecuador: Napo 
Region, SE of Coca, near Rio Tiputini, Yasuni National 
Park, 260 m, 15-25.i.2000 (Puplesis & Hill), genitalia 
slide no. 29612 (BMNH). Excluded from paratype 
status: 1 ?, data as holotype, genitalia slide no. AD0330 

Remarks. A single female has been associated with 
the holotype since it has very similar dark forewings 
with fuscous apical irroration; it is similarly small and 
was collected at the same time in the same locality. 
However, there are some points of difference which 
make us preclude this specimen from paratype status: 
1 ) the palpi are whitish, not yellowish cream as in the 
male; 2) the head is slightly rubbed and comparison 
with the holotype is difficult, but the tuft still present is 
quite whitish (not orange-ochreous as in the male); 3) 
there is no pale postmedial area developed on the 
forewing as in the male; 4) the abdomen is silvery on 
the underside, not brown. 

Acalyptris insolentis Puplesis & Diskus sp. n. 

(Figs 19,83-88) 

MALE (Fig. 1 9). Forewing length: 1 .7-2.0 mm. Wing- 
span: 3.8^.5 mm. Head: palpi cream; frontal tuft 
orange or pale ochreous; collar indistinct, comprised 
of cream piliform scales; eyecaps cream to whitish; 
antenna brownish, comprising of about 24 segments. 
Thorax and tegulae yellowish cream. Forewing with 
slightly variable but distinct pattern; basal area before 
fascia, except large costal area, covered with small, 
pale brownish scales; costal area distinctly paler, 
yellowish cream, as tegulae; fascia slightly postmedial, 
distinctly oblique and broadening at tornus. formed by 
fuscous brown and brown scales; area beyond the 
fascia yellowish cream; apex of forewing with yellow- 
ish cream background but irrorated with brown and 
fuscous scales; some fuscous scales overlapping onto 
cilia. Cilia cream only at distal end of forewing, 
otherwise grey or greyish. Underside of forewing brown 
to ochreous brownish. Hindwing lanceolate, brown or 
pale brownish depending upon angle of observation; 
cilia greyish to grey. No androconial patches on 
hindwing or forewing. Legs ochreous cream. Abdomen 
brown on upperside, yellowish brown on underside, 
with a pair of cream abdominal tufts composed of long 
piliform scales. 

Female. Unknown. 
Genitalia c? (Figs 83- 

Capsule 270-280 ^m 

long. Tegumen extended into short, broadly rounded 
pseuduncus. Uncus (Fig. 84) with lateral, well- 
sclerotized, rod-like parts and broadened, lobe-like 
caudal element. Gnathos (Fig. 85) complex and dis- 
tinctive; central plate large, with oval anterior extension 
and broad lobe-like caudal extension homologous with 
caudal process of other Acalyptris; the last weakly 
sclerotized, almost membranous, usually inconspicu- 
ous even in stained slides; lateral arms of vinculum 
very broad, almost as broad as long. Tegumen with two 
distinct, lateral rod-like sclerites along capsule beneath 
valvae. Valva 1 76 u.m long, straight, generally narrow, 
slightly bulged inwardly before apex and at base; no 
processes developed. Transtilla with very slender trans- 
verse sublateral processes, more than one-half length 
of transverse bar which may be straight in contrast to 
Fig. 83. Juxta formed by vinculum posterior margin 
and attached to ventral side of aedeagus, a small, 
sclerotized, variable, oval to rounded, triangular or 
square lobe between bases of valvae. Vinculum very 
short with unusually long, slender, distally truncate 
lateral lobes; anterior excavation square and deep. 
Aedeagus 212-220 urn long, with two large paired 
lateral carinae and large, ventral lobe-like extension 
attached to vinculum: cornuti either absent, or occasion- 
ally represented by an indistinct single spine. 

Biology. Adults collected in January. 

DIAGNOSIS. Externally this species resembles the 
Central American hispidus Puplesis & Robinson. 
novenarius Puplesis & Robinson and laxibasis Puplesis 
& Robinson; however insolentis differs in the presence 
of a large, yellowish cream costal area. In the male 
genitalia the characteristic gnathos with caudal and 
anterior extensions distinguish this species from all 
known Acalyptris; the lobate caudal component of the 
uncus together with the slender lateral lobes of the 
vinculum are also, in combination, distinguishing 

Distribution. Premontane and lowland Amazon 
rainforest (Ecuador); thus apparently more widespread, 
at least in the western Amazon basin, than any other 
species found so far. 

Condition of type material. The holotype and 
three paratypes are in good condition but the frontal 
tuft of the holotype is rubbed; the fourth paratype 
( AD0323 VPU) has the detached hindwings stored in a 
small gelatine capsule beneath the specimen. 

Material examined. Holotype d\ Ecuador: Napo 
Region: near Rio Napo, Jatun Sacha, premontane rain- 
forest, 500 m, 26-3 1 .i.2000 (Puplesis & Hill), genitalia 
slide no. 29621 (BMNH). Paratypes: 2 c?, data as 
holotype, genitalia slides nos 29622 (BMNH) and 
AD0329 (VPU); 1 c?, SE of Coca, near Rio Tiputini, 
Yasuni National Park, 260 m, 15-25.i.2000 (Puplesis 
& Hill), genitalia slide no. AD0323 (VPU). 





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de la sociedad Entomologica Argentina. Buenos Aires, 23: 3 1 — 46. 

Davis, D.R. 1978. New leaf-mining moths of the family Nepticulidae 
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Neotropical Lepidoptera. Checklist: part I. Micropterigoidea - 
hnmoidea. Junk, den Haag. xxvii + 1 1 2 pp. 

Davis, D.R. 1 985. A re-examination of Enleucha cyanochlora Meyrick 
and its subsequent transfer to the Nepticulidae (Lepidoptera: 
Nepticuloidea). Proceedings of the Entomological Society of Wash- 
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Forbes, W.T.M. & Leonard, M.D. 1 930. A new leaf-miner of cotton 
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Meyrick, E. 1 9 1 5. Descriptions of South American Micro- Lepidoptera. 
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36: 377-415. 

Newton, P.J. & Wilkinson, C. 1982. A taxonomic revision of the 
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Principal references are in bold. 

acuta 21 
amazonius 32 
andina 23 
articulosus 30 
austroamericana 25 
barbata 26 

basihastatus 29. 30. 31 
cuprata 23 
diskusi 27 
dividua 28. 29 
ecuadoriana 27, 29 



gilvafascia 21 
guajavae 22 
hispidus 32. 33 
insolentis 33 
latipennata 28, 29 
laxibasis 33 
martinheringi 29,30,31 
molybditis 27 
montanotropica 23 
novenarius 32, 33 
nubimontana 24. 25 
onorei 28 
platygnathos 32 
plumosetaeella 26 
pseudohastatus 30. 31 
repanda 26 
rotundus 31 
rubeta 24, 25 
salicis group 23, 24 
snaddoni 21 
species 29122 27 

tabulosa 27 



Figs 1-3. Neotropical Nepticulidae. 1 , Acalyptris onorei forewing venation; 2. Enteucha guajavae forewing venation; 3, 
same, adult moth. 



,: • t$Ml Vi ^ 

Figs 4—7. Neotropical Nepticulidae. 4, Enteucha acuta: 5, Stigmella montanotropica; 6, S. nubimontana; 7, 5. rubeta. 





Figs 8-11. Neotropical Nepticulidae. 8, Stigmella austroamericana; 9, Fomoria repanda; 10, F. tabulosa; 1 1 , Acalyptris 



>' - mm 

lie V « 




Figs 12-15. Acalyptris species. 12, onorei; 13, same, hindwing; 14, basihastatus; \5,pseudohastatus. 







Figs 16-19. Acalyptris species. 16, articulosus; 17, rotundas; 18, amazonius; 19, insolentis. 



Figs 20-25. Male genitalia of Enteucha guajavae. 20, holotype (AD0340-VPU) uncus; 21, same, capsule; 22, same, 
aedeagus; 23, paratype (AD0342-VPU) aedeagus; 24, paratype (AD0341-VPU) transtilla; 25, same, aedeagus. Scale: 
0.1 mm. 



Figs 26-30. Male genitalia of Nepticulidae. 26, Enteucka acuta, holotype (29604-BMNH) capsule; 27, same, aedeagus, 28, 
same, right valva and juxta; 29, Stigmella montanotropica, paratype, (29618-BMNH) aedeagus; 30, same, capsule. Scale: ' 

0.1 mm. 






Figs 31-36. Male genitalia of Stigmella nubimontana. 31, holotype (29614-BMNH) capsule; 32, same, aedeagus; 33, 
paratype (AD0322-VPU) uncus; 34, paratype (AD0321-VPU) uncus; 35, paratype (AD0321-VPU) apex of valva; 36, 
paratype (AD0322-VPU) both valvae and transtilla. Scale; 0.1 mm. 



Figs 37-43. Male genitalia of Stigmella. 37, rubeta, holotype (29616-BMNH) capsule; 38, same, tegumen + uncus; 39, 
same, aedeagus; 40, austroamericana, holotype (29613-BMNH) uncus; 41, same, valva; 42, same, aedeagus; 43, same, 
capsule. Scale: 0.1 mm. 



Figs 44-48. Male genitalia of Fomoria repanda, holotype (29603-BMNH). 44, capsule; 45, aedeagus; 46, valva, 47, 
pseuduncus; 48, gnathos. Scale: 0.1 mm. 



Figs 49-52. Male genitalia of Fomoria repanda. paratype (AD0315-VPU). 49. capsule; 50. pseuduncus; 51, uncus + 
gnathos: 52, aedeagus. Scale: 0.1 mm. 




Figs 53-58. Male genitalia of Nepticulidae. 53, Fomoria tabulosa, holotype (29625-BMNH) capsule; 54, same, juxta; 55, 
same, aedeagus; 56, Acalyptris ecuadoriana, holotype (29624-BMNH) capsule; 57, same, gnathos; 58, same, aedeagus. 
Scale: 0.1 mm. 



Figs 59-64. Male genitalia of Acalyptris onorei, holotype (29623-BMNH). 59, capsule; 60, valva; 61, pseuduncus; 62, 
paired uncus; 63. paired gnathos; 64, aedeagus. Scale: 0.1 mm. 



Figs 65-67. Male genitalia of Acalyptris basihastatus. 65, holotype (29605-BMNH) capsule; 66. paratype, (29606-BMNH) 
pseuduncus; 67, holotype (29605-BMNH) aedeagus. Scale: 0.1 mm. 



Figs 68-70. Male genitalia of Acalyptris pseudohastatus, holotype (29607-BMNH). 68, capsule; 69, pseuduncus; 70, 
aedeagus. Scale: 0.1 mm. 



Figs 71, 72. Valvae of Acalyptris. 7 1 , basihastatus, paratype (29606-BMNH); 72, pseudohastatus, holotype (29607- 
BMNH). Scale: 0.1 mm. 



Figs 73-78. Male genitalia of Acalyptris. 73, articulosus, holotype (29610-BMNH) capsule; 74, same, pseuduncus; 75, 
same, aedeagus; 76. rotundus, holotype (2961 1-BMNH) capsule; 77, same, pseuduncus; 78, same, aedeagus. Scale: 
0.1 mm. 



Figs 79-88. Male genitalia of Acalyptris. 79, amazonius, holotype (29612-BMNH) capsule; 80, same, pseuduncus; 81, 
same, uncus; 82, same, aedeagus; 83, insolentis, holotype (29621-BMNH) capsule; 84, same, uncus; 85, same, gnathos; 
same, pseuduncus; 87. same, aedeagus with broken ventral lobe (juxta); 88. paratype (29622-BMNH) aedeagus. Scale: 
0.1 mm. 



Figs 89, 90. Female genitalia of Enteucha guajavae. 89, paratype (AD0344-VPU); 90, paratype ( AD0343-VPU). Scale: 
0.1 mm. 



Fig. 91. Female genitalia of Stigmella montanotropica, holotype (296 1 9-BMNH). Scale: 0. 1 mm. 



Figs 92, 93. Female genitalia of Nepticulidae. 92, Stigmella ntbeta, paratype (296 1 7-BMNH); 93, Acalyptris amazonius, 
specimen excluded from type series (AD0330-VPU). Scale: 0.1 mm. 







■ ,..; r.*/ 


Figs 94-97. Mines of Stigmella species on Rubus spp. from Ecuador. 94, nubimontana, damaged leaf with a few mines, 
high Andes, Papallacta, sample no. 4630-VPU; 95, same, combined mine (gallery + blotch) showing frass distribution; 96, 
rubeta, damaged leaf with two mines, tropical western slopes of Andes, Tandapi, sample no. 4629- VPU; 97, same, 
combined mine (gallery + blotch) showing frass distribution. 





Ifl§; ^f£| 





Figs 98-100. Mines of Nepticulidae. 98, Stigmella montanotropica, damaged leaf of Acalypha with a sinuous mine, western 
slopes of Andes, Tandapi, sample no. 4628-VPU; 99, same, gallery showing frass distribution; 100. Enteucha guajavae, 
damaged leaf of Psidium guajava with mines, western foothills of Andes, Bucay, sample no. 4722- VPU. 





Figs 101, 102. Habitats of species. 101 , Enteucha guajavae, western foothills of Andes, 700 m Bucay (=Cumanda); 102, 
Stigmella montanotropica, western slopes of Andes, 1200 m, Tandapi. 

X^C 3 ^^° 2 (1 J 

ft///. w/r Hist. Mus. Lund. (Ent.) 71(1): 59-76 Issued 27 June 2002 

A review and checklist of the 
Neotropical Nepticulidae 


Department of Zoology, Vilnius Pedagogical University, 39, Studentu str., Vilnius 
2004 LT, Lithuania 


Department of Zoology, Vilnius Pedagogical University, 39, Studentu str., Vilnius 
2004 LT, Lithuania 


Department of Entomology, The Natural History Museum, Cromwell Road, London 


Departamento de Biologia, Pontificia Universidad Catdlica del Ecuador, Quito, 


Synopsis 59 

Introduction 60 

Material and methods 60 

Abbreviations of institutions 61 

Acknowledgements 61 

Redescriptions of Manoneura basidactyla and Ectoedemiafuscivittata 61 

Revised checklist of Neotropical Nepticulidae 63 

Discussion 63 

Definition of species-groups, taxonomic notes 63 

Leaf-mine collecting and hostplant data in equatorial America 66 

Diversity and geography 66 

References 67 

SYNOPSIS. Fieldwork in 2000-200 1 has added substantially to our knowledge of the Nepticulidae 
of the upper Amazon basin and the Andes and increased the number of species known from the 
Neotropical Region from 58 to 74. Two species - Manoneura basidactyla (Davis) and Ectoedemia 
fuscivittata Puplesis & Robinson -are recorded from equatorial America for the first time here and 
are redescribed, with amplified descriptions and illustrations. A revised checklist of the Neotropical 
Nepticulidae is given together with an updated distribution map for Central and Southern America. 
Fomoria latipennata Puplesis & Robinson is transferred to Acalyptris, comb. n. Four new species- 
groups (Stigmella tiliella-group, S. barbata-group, Fomoria molybditis-group and Acalyptris 
latipennata- group) are defined. Hostplant data are reviewed and further hostplant genera from 
which mines or cocoons have been collected are noted. The diversity and distribution of the 
Neotropical Nepticulidae are discussed. 

© The Natural History Museum. 2002 





Nepticulidae are a family of minute monotrysian 
Microlepidoptera with a worldwide distribution and 
about 750 described species. Their morphology, 
biology and taxonomic composition have been recently 
reviewed (Puplesis & Robinson, 2000). 

The size of the adults, concealed mining life-style 
of the larvae (predominantly in leaves), and the diffi- 
culty of rearing imagines goes some way towards 
explaining why these moths are still poorly studied in 
many regions. Only the northern European nepticulid 
fauna can be considered to have been exhaustively 
studied. Studies in other regions of the world in all 
probability underestimate the diversity of the group. 
Key works are reviewed by Puplesis & Robinson 

The history of description of Nepticulidae from 
the Neotropical Region has been reviewed by 
Puplesis & Robinson (2000). That paper, based on 
the results of fieldwork in Belize in 1998 by Puplesis 
and Simon Hill (UW) together with investigation 
of unidentified material in ZMUC and USNM, 
documented a total of seven genera and 58 species 
of Nepticulidae from Central and Southern America. 
Twenty-eight of those species recognized were 
new taxa from Belize (including four species left 
unnamed). None was from the "heart" of the Neotropics 
- the Amazon basin - and it was assumed that 
the absence of specimens from Amazonian rainforest 
reflected collecting effort, coupled with a lack 
of diversity. 

In the preceding paper in this journal (Puplesis, 
Diskus & Robinson, 2002) sixteen new species from 
the upper Amazon basin and the Andes (Ecuador) are 
recorded, increasing the number of species known 
from the neotropics by more than one-fifth. The 
fieldwork in Ecuador upon which that paper is based 
resulted in the acquisition of additional material of 
Manoneura basidactyla (Davis) and Ectoedemia 
fuscivittata Puplesis & Robinson. These are the first 
records of these taxa from equatorial America. In this 
paper we amplify the descriptions and provide further 
illustrations of these species and update the checklist 
of Neotropical Nepticulidae with a distribution chart 
and map. We define four species-groups and review 
the known biologies of neotropical Nepticulidae, add- 
ing observations on nepticulid leaf-mines in Ecuador, 
and review the diversity of Nepticulidae in tropical 

The present collaborative project was undertaken at 
the Natural History Museum, London with the support 
of the Royal Society (London), Professor Hering 
Memorial Fund (London) and in cooperation with the 
Pontificia Universidad Catolica del Ecuador, Quito, 
Ecuador (Dr Giovanni Onore). 

Brief details of collecting localities in Ecuador (Fig. 1) 
were given in the preceding paper (Puplesis, Diskus & 
Robinson, 2002 - q.v.). Primary sites were at the 
Yasuni Research Station, Jatun Sacha Biological Sta- 
tion and Misahualli (Figs 2-4). 

Jatun Sacha and the nearby village of Misahualli 
(8 km NW of Jatun Sacha) are located on the south side 
and the northern side of the upper Rio Napo respec- 
tively, about 20 km east of the base of the Andes. The 
environment is transitional between the lower slopes 
of the Andes and Amazon lowland. The area is formed 
by steep hills crossed by seasonal streams. The soil is 
mostly a red clay oxisol. Annual precipitation is 
3900 mm, fairly evenly distributed throughout the year, 
though the collecting periods (January 2000 and Janu- 
ary 2001) tended to be relatively dry. Around half of 
Jatun Sacha Reserve is covered with original undis- 
turbed forest, while the remainder is secondary growth; 
Misahualli (Fig. 4) is mostly surrounded by secondary 
forest and only about 10 to 15% can be considered as 
primary forest. The most common plants in the area 
are: Adiantum cayennense Klotzsch, Asplenium 
auritum Swartz, Anthurium balslevii Croat, Bactris 
gasipaes H.B.K., Guzmania acuminata L.B.Smith, 
Becquerelia cymosa Brongn, Calathea attenuata 
Kennedy, Heliconia stricta Huber, Epidendrum 
coronation Ruiz & Pavon, Maxillaria tarumanensis 
Hoehne, Schefflera diplodactyla Harms and Matelea 
rivularis Woodson. 

Yasuni Research Station is based at the Yasuni 
National Park and Biosphere Reserve, which together 
cover approximately 9820 square kilometres of mainly 
pristine Amazon lowland rain forest, rivers, lagoons 
and swamps (Figs 2, 3). The annual precipitation pattern 
is very similar to Jatun Sacha. The most common 
plants in the area are: Alseis lugonis (L.) Andersson, 
Astrocaryum urostachys Burret, Guatteha glaberrima 
(R.E.) Fr., Trigynaea triplinervis D.M. Johnson & 
N.A. Murray, Nectandra crassdoba Rohwer, Euterpe 
precatoria Mart., Ceiba samauma (Mart.) K. Schum, 
Phragmotheca ecuadorensis W.S. Alverson, Duguetia 
spixiana Mart., Oxandra mediocris Diels, Aristolochia 
goudotti Duch., Mauritia flexuosa L.F., Ceiba 
pentandra (L.) Gaertn., Inga tessmannii Harms and 
Miconia grandifolia Ule. 

Collecting methods, techniques for genitalia 
preparation and protocols for description are outlined 
in the preceding paper (Puplesis, Diskus & Robinson, 
2002 - q.v.). Black and white drawings cannot show 
the metallic lustre (especially of the lustrous 
Manoneura species) and the coloured iridescence 
characteristic of most Nepticulidae, and details of such 
colours have been incorporated into species descrip- 



Genitalia drawings were made by Puplesis using a 
camera lucida from permanent slides. 


BMNH The Natural History Museum, London, UK 
(formerly British Museum (Natural History)) 

VPU Vilnius Pedagogical University. Vilnius, 


USNM National Museum of Natural History. Washing- 
ton DC, USA (formerly United States National 

ZMUC Zoologisk Museum, University of Copenhagen, 
Copenhagen, Denmark 

Acknowledgements. See the precedi ng paper ( Puplesis, 
Diskus & Robinson. 2002) for relevant acknowledgements 
for assistance, funding and collecting and study permissions. 

We are grateful to Mr Arturas Skorupskas (VPU) for making 
indian-ink drawings of the adults; Mrs Birute Noreikiene 
(VPU) is thanked for making the drawing of the adult of 
Ectoedemiafuscivittata from Belize. 

Arunas Diskus thanks The Natural History Museum for 
financial support for studies conducted under the SYS- 
RESOURCE Programme (project manager Vanessa Pike). 


Manoneura basidactyla (Davis, 1978) 

(Figs 6, 9-15, 21) 

Oligoneura basidactyla Davis, 1978: 218-219. 
Manoneura basidactyla Davis: Puplesis & Robinson, 
2000: 22,23, figs 12, 61, 83-85, 207. 

Male (Fig. 6). Forewing length: 1 .7-2. 1 mm (Cen- 
tral American specimens) and about 2.2 mm 
(Amazonian specimens). Wingspan: 4.3-4.9 mm. 
Head: palpi yellowish cream to ochreous cream or 
cream; frontal tuft orange; collar a large tuft of lamel- 
lar yellowish cream scales with golden reflection; 
eye-caps yellowish cream; antenna brownish grey, ca. 
38^44 segments. Thorax, tegulae and forewing fuscous 
brown with bronze or copper (in Amazonian specimens) 
iridescence and very strong blue and purple reflections. 
Distinct postmedian fascia of forewing oblique, 
yellowish gold. Cilia fuscous, tending be lighter distally. 
Underside of forewing brown to dark brown (with 
purple and bluish iridescence in Amazonian 

specimens). Hindwing lanceolate, very slender, dark 
brown with strong purple and blue reflections which 
are stronger on upperside of hindwing; cilia brownish. 
No androconia on forewing or hindwing. Legs grey to 
fuscous but tarsi cream. Abdomen black on upperside, 
blackish grey on underside; genital segments blackish 
grey, not contrasting with main colour of abdomen. 

Female. Similar to male. Forewing length: 2.2-2.4 
mm. Wingspan: 5.3-5.4 mm. Antenna ca. 32-38 seg- 
ments. Legs cream with black lateral shading or mainly 
fuscous (in Amazonian specimens). Abdomen black- 
ish on upperside but cream or creamy brown on 
underside. Otherwise as male. 

Genitalia d (Figs 9-14). Capsule ca. 290-305 |im 
long. Tegumen with short caudally slightly bilobed 
pseuduncus-like extension. Uncus with strong pointed 
central process directed anteriorly, and long narrow 
lateral arms directed posteriorly. Gnathos with small 
but complex and well-sclerotized v-shaped central 
region and rather membranous broad lateral arms; 
shape and sclerotization of central element may vary. 
Natural position of central pointed process of uncus 
very closely appressed to posterior excavation of central 
part of gnathos (appearing to be a single structure) 
(Fig. 1 0). Valva ca. 205-2 1 5 (am long, relatively narrow 
in distal half and gradually broadened towards base; 
apical process long and slender. Transtilla absent, i.e., 
no transverse bar; bases of valvae with remarkably 
long and straight apodemes. Vinculum very long and 
very broad, truncate at anterior end; no anterior 
excavation or lateral lobes. Aedeagus 236-246 u.m 
long, with two pairs of pointed lateral carinae: no 
cornuti on vesica. Juxta a band-like sclerite, abruptly 
broadened basally, fused with aedeagus. 

Genitalia 9 (Fig. 15). Total lengthca. 720-915 fim. 
Anal papillae undeveloped. S8 and T8 widely rounded. 
Apophyses posteriores short and complex. Apophyses 
anteriores very slender, 0.5-0.7 length of apophyses 
posteriores. Vestibulum sclerotized. Caudal part of 
corpus bursae very narrow; remaining part ovally broad- 
ened; no signa visible. Accessory sac undeveloped, in 
Central American specimens represented by a small 
but clearly visible ring-like sclerotization; ductus 
spermathecae always long and narrow, slightly sinuous. 

Biology. Hostplant: Coccoloba uvifera (L.) L., 
Polygonaceae (for the Caribbean fauna; specimens 
from Amazon rainforest have not been reared and the 
hostplant is not established). Adults collected in Janu- 
ary, April-May and July. The suggestion that the species 
might be univoltine (Davis, 1978) is not supported. 

DIAGNOSIS. M. basidactyla may be distinguished 
from the other species of the genus (trinaria) by the 
relatively straight valva and distally truncate vincu- 



lum; externally it is similar in pattern and iridescence, 
but basidactyla is slightly paler. It differs from other 
nepticulids from the Neotropical Region (except 
Manoneura trinaria) in the strong purple reflection of 
the forewing and the large distally truncate vinculum 
together with the very specialized shape of the gnathos 
(shared only with trinaria). 

Distribution (Fig. 21). Southwest coast of Florida, 
Dominica, Belize, Ecuador. It is likely that the species 
has a wide distribution in tropical forest in the 

Material examined. Belize: Chiquibul Forest 
Reserve, Las Cuevas, 3 d\ 2 9, 3-16.iv. 1998 (Puplesis 
& Hill) genitalia slide no. 29120d\ no. 29121? 
(BMNH); 2c?, 1 9, same data, wing venation slide no. 
AD0314 9 (VPU). Dominica: Pont Casse, Id", 
16.V.1965 (Davis) genitalia slide no. Diskus 002 
(USNM);CabritSwam, 1 d\ 1 9, 10-13.v.l965(Dav/s) 
(USNM); Springfield Est., 2 specimen (no abdomens), 
20-26.vii.1963 {Flint) (USNM). Ecuador: Napo 
Region, SE of Coca, near Rio Tiputini, Yasuni National 
Park, 260 m, 1 d\ genitalia slide no. AD0327 VPU, 
1 9, genitalia slide no. AD0326 VPU, 15-25.i.2000 
(Puplesis & Hill). 

Ectoedemia fuscivittata Puplesis & Robinson, 

(Figs 7, 8, 16-21) 

Ectoedemia fuscivittata Puplesis & Robinson, 2000: 
42, figs 39, 154-156,223. 

Male (Figs 7, 8). Forewing length: 1.7-1.8 mm. 
Wingspan: 4. 1^4.3 mm. Head: palpi cream to ochre - 
ous cream; frontal tuft pale ochreous to pale 
orange-ochreous; collar indistinct, pale ochreous, 
comprised of piliform scales; eyecaps ochreous cream, 
moderately large; antenna brownish or greyish to pale 
brownish ochreous, 42^44 segments. Thorax anteriorly 
ochreous yellow, elsewhere metallic grey; tegulae 
ochreous yellowish with few greyish scales anteriorly. 
Forewing basal three-fifths to fascia metallic grey to 
ochreous cream (Amazonian specimen) with or with- 
out (Amazonian specimen) distinctive blue-green and 
some indistinct purplish reflection; entire forewing of 
Amazonian specimen, or, in Belize specimens, narrow 
area along costa, particularly before fascia and on 
tornus, and area beyond fascia with ochreous cream 
scales; distinct postmedian fascia oblique, fuscous 
brown, but weakly defined in Amazonian specimen; 
with some dark brown scales in area before cilia. Cilia 
metallic greyish or dark ochreous cream in Amazonian 
specimen. Underside of forewing grey-brown orbrown- 
ochreous. Hindwing pale brownish, cilia greyish to 
brownish grey; a long patch of whitish scales may be 

visible on basal half of hindwing upperside, however 
these androconia not always distinctive. Legs cream or 
ochreous, with or without (Amazonian specimen) grey- 
fuscous lateral shading. Abdomen fuscous brown on 
upperside, brown or brownish on underside; genital 
segments mostly covered by dark (predominantly 
brown) scales, not contrasting with main colour of 

Female. Unknown. 

Genitalia S (Figs 16-20). Capsule 268-270 urn 
long. Pseuduncus small, distinctly rounded and strongly 
papillated. Dorsal plate of tegumen simple, small. 
Gnathos with triangular caudal process, slender lateral 
arms, and small oval central plate. Valva 150-170 u.m 
long, distinctly triangular, with more or less straight 
inner margin tapering into pointed, caudally directed 
apical process; in Amazonian specimen apical 
processes particularly narrow (Fig. 19). Basal margins 
of valva strongly sclerotized. Transtilla without 
transverse bar, but with rather long and slender or very 
slender valval apodemes. Juxta absent, valvae fused 
via basal membranous joint. Vinculum very small, 
with very small but distinctly shaped and well- 
sclerotized lateral lobes; in Amazonian specimen the 
lobes broader. Anterior excavation of vinculum always 
very shallow, broad or narrow (Amazonian specimen). 
Aedeagus 210-235 um long, abruptly bulged in basal 
1/2-2/3 or gradually broadened (Amazonian speci- 
men); without carinae and weakly sclerotized apical 
part; vesica with some very tiny indistinct cornuti only; 
strong sclerotization of aedeagal tube in Amazon speci- 
men may appear as a long spine-like cornutus but it is 
not such (genitalia slide no. AD0328 VPU, Fig. 20). 

Biology. Adults collected in April (Belize) and Janu- 
ary (Ecuador). 

Diagnosis. Although many species belonging to 
Ectoedemia are not easily distinguishable from each 
other, fuscivittata is an exception: the combination of 
features such as the caudally directed valval process, 
basally broadened aedeagus, very short vinculum, and 
dark oblique forewing fascia make this a very distinc- 
tive species. The absence of a transverse transtilla bar 
in this species sets it apart from all other Ectoedemia. 

Distribution (Fig. 21). 

Rainforest in Belize and 

Material examined. Belize: Cayo District, 
Chiquibul Forest Reserve, Las Cuevas, 3 c? [holotype 
(BMNH) and paratypes (BMNH, VPU)] 3-16.iv. 1 998 
(Puplesis & Hill) genitalia slides no. 29107 [holotype], 
no. AD 0302 [paratype, VPU]. Ecuador: Napo Re- 
gion, SE of Coca, near Rio Tiputini, Yasuni National 
Park, 260 m, 1 c?, genitalia slide no. AD0328 VPU, 15- 
25.i.2000 (Puplesis & Hill). 




Seventy-four species are now recognized from the 
Neotropical Region (including most of Mexico, 
together with the US states of Arizona and Florida) and 
these are listed in Table 1 . Another eight species with a 
distribution range from the north of Florida into the 
northern states of the USA should not be considered to 
be part of the neotropical fauna; Stigmella 
nigriverticella (Chambers), S. castaneaefoliella 
(Chambers), S. ostryaefoliella (Clemens), 
S. myricafoliella (Busk), Ectoedemia clemensella 
(Chambers), E. similella (Braun), E. virgulae (Braun), 
E. obrutella (Zeller) have predominantly boreal distri- 
bution ranges and/or hostplants. 


Definition of species-groups, taxonomic 

Venation. In studying neotropical Nepticulidae we 
have met difficulties in using wing venation to clarify 
the generic position of species. Although wing venation 
can provide useful characters for generic diagnosis of 
many Holarctic or South African Nepticulidae (Scoble, 
1983; Nieukerken, 1986; Puplesis, 1994), a few 
Neotropical species show unusually strong reduction 
of the venation, limiting the wider value of venational 
features. This is notable in some Fomoria such as 
F. diskusi (Puplesis & Robinson, 2000: fig. 64). On the 
other hand, a few other neotropical species have a less 
derived venation than Old World members of the same 
genus. In the forewing of Enteitcha guajavae, Stigmella 
ovata andAcalyptris onorei R4 and R5 are represented 
by two separate veins and are not coalescent as is usual 
in these genera (see Puplesis & Robinson, 2000: fig. 
62; Puplesis, Diskus & Robinson, 2002: figs 1,2). The 
accepted generic diagnoses of these groups are based 
upon the Old World taxa and need revision in the light 
of the morphology of the neotropical representatives. 

Stigmella salicis-group. This long-established group 
contains more than 30 described species worldwide. It 
was first designated for European species and currently 
contains 15 described Palaearctic representatives. How- 
ever, it also has at least two representatives in the 
Nearctic (known as the \fuscotibiella-group' ). Puplesis 
& Robinson (2000) recognized 14 mostly Andean 
species as belonging here. All Holarctic species (except 
one on Vaccinium) feed on Salicaceae (Salix, Populus) 
while Neotropical taxa feed on Rosaceae (Rubas) and 
Euphorbiaceae (Acalypha). The female genitalia of 

most of the Neotropical species lack the characteristic 
signum of tiny, dentate chitin plates encircling the 
bursa that is a derived feature and typical of the 
Holarctic members of the group; just olyritis and 
montanotropica (see Puplesis, Diskus & Robinson, 
2002: Fig. 91) have a similar structure on the bursa. 
The male genitalia of the Neotropical species are simi- 
lar to those of Holarctic salicis-group species. The 
Holarctic representatives may form a monophyletic 
entity within the salicis-group, defined by the presence 
of a signum, and derived from a Neotropical-type 

Stigmella tiliella-group. A new species-group is 
designated here for two closely related and distinctive 
species. A rounded valva with a short dorsal process, 
broadly U-shaped gnathos and a cluster (or clusters) of 
spine-like cornuti in the aedeagus are characteristic of 
the male genitalia. The group resembles the Holarctic 
paradoxa-group but differs in the cornuti clusters and 
gallery mines (which in the paradoxa-group are com- 
bined, i.e., distinctly blotch-shaped in the second half 
of the course). The group contains Stigmella tiliella 
(Braun) known from the Nearctic (USA, Kentucky) 
and S. kimae Puplesis & Robinson, 2000 (Belize). The 
first makes gallery-type mines on Tilia americana 
leaves; the hostplant of the second species is still 

Stigmella barbata-group. The group is newly 
designated here for species possessing unique plumose 
scales on the apex (or apical third) of the valva; in 
contrast to most Stigmella, the complement of cornuti 
in the aedeagus is very weakly developed, the cornuti 
tiny and weakly sclerotized. The group appears to be 
endemic to the Neotropics and contains three species: 
Stigmella plumosetaeella Newton & Wilkinson. 1982, 
barbata Puplesis & Robinson, 2000 and 
austroamericana Puplesis & Diskus, 2002; the biol- 
ogy of these species is unknown. The recently 
discovered Amazonian austroamericana appears to be 
the sister-group of the Central American barbata and 
the two appear to represent an allopatric and vicariant 

Fomoria. Although some species of Fomoria may 
be clearly grouped as monophyletic units, the taxo- 
nomic status of the genus itself is questionable because 
its monophyly is unproven. The genitalia of Fomoria 
and Ectoedemia (sensu stricto) follow the same ground 
plan, but the uncus (which is entirely reduced in 
Ectoedemia) is fully preserved in Fomoria (a 
plesiomorphy). Vein Cu in the forewing tends to be 
shortened or completely lacking (apomorphy) in 
Fomoria, but this character is not always consistent, at 
least among the Neotropical representatives (see above: 


Table 1. Distribution of Neotropical Nepticulidae. The USA column includes only species from Arizona and Florida which 
are recognized as tropical; 'boreal' species with northern distribution ranges and/or temperate hostplants are excluded. 

No Species 

— c 

ENTEVCHA Meyrick, 1915 

1 cyanochlora Meyrick, 1915 

2 gilvafascia (Davis. 1978) 

3 hilli Puplesis & Robinson, 2000 

4 contracolorea Puplesis & Robinson, 2000 

5 terricula Puplesis & Robinson, 2000 

6 snaddoni Puplesis & Robinson. 2000 

7 guajavae Puplesis & Diskus, 2002 

8 acuta Puplesis & Diskus, 2002 

MANONEURA Davis, 1979 

9 hasidactyla (Davis, 1978) 

10 trinaria Puplesis & Robinson, 2000 
STIGMELLA Schrank, 1802 
The salicis species-group 

11 andina (Meyrick, 1915) 

1 2 cuprata (Meyrick. 1915) 

13 johannis (Zelller, 1877) 

14 rudis Puplesis & Robinson, 2000 

1 5 marmorea Puplesis & Robinson, 2000 

16 peruanica Puplesis & Robinson, 2000 

17 epicosma (Meyrick, 1915) 

18 schoorli Puplesis & Robinson, 2000 

19 hamata Puplesis & Robinson. 2000 

20 imperatoria Puplesis & Robinson. 2000 

21 olyritis (Meyrick, 1915) 

22 montanotropica Puplesis & Diskus, 2002 

23 nubimontana Puplesis & Diskus, 2002 

24 rubeta Puplesis & Diskus, 2002 
The eurydesma species-group 

25 eurydesma (Meyrick. 1915) 

26 albilamina Puplesis & Robinson, 2000 

27 fuscilamina Puplesis & Robinson, 2000 
The tiliella species-group 

28 kimae Puplesis & Robinson, 2000 
The barbata species-group 

29 plumosetaeella Newton & Wilkinson, 1982 

30 barbata Puplesis & Robinson, 2000 

3 1 austroamericana Puplesis & Diskus, 2002 
Unattributed to a species-group 

32 gossypii (Forbes & Leonard. 1930) 

33 pruinosa Puplesis & Robinson, 2000 

34 ovata Puplesis & Robinson, 2000 

35 hylomaga (Meyrick, 1931) 

36 costalimai (Bourquin, 1962) 

37 guittonae (Bourquin, 1962) 

Table 1 continued 



ECTOEDEMIA Busck, 1907 
reneella Wilkinson, 1981 
helenella Wilkinson, 1981 
mesoloba Davis. 1978 
species 29105 

fuscivittata Puplesis & Robinson, 2000 
FOMORIA Beirne, 1945 

tabulosa Puplesis & Diskus, 2002 

The molybditis species-group 

molybditis (Zeller, 1877) 

diskusi Puplesis & Robinson, 2000 

species 29122 

repanda Puplesis & Diskus. 2002 

ACALYPTRIS Meyrick, 1921 
The latipennata species-group 

latipennata (Puplesis & Robinson. 2000) comh.n. 

dividua Puplesis & Robinson. 2000 

ecuadoriana Puplesis & Diskus, 2002 

onorei Puplesis & Diskus. 2002 

Unattributed to a species-group 

bovicorneus Puplesis & Robinson. 2000 

martinheringi Puplesis & Robinson. 2000 

fortis Puplesis & Robinson, 2000 

hispidus Puplesis & Robinson. 2000 

novenarius Puplesis & Robinson. 2000 

lascuevella Puplesis & Robinson, 2000 

bifldus Puplesis & Robinson, 2000 

trifldus Puplesis & Robinson. 2000 

tenuijuxtus (Davis. 1978) 

unicornis Puplesis & Robinson, 2000 

laxibasis Puplesis & Robinson, 2000 

bicornutus (Davis, 1978) 

species 29135 

platygnathos Puplesis & Robinson. 2000 

species 29140 

basihastatus Puplesis & Diskus, 2002 

pseudohastatus Puplesis & Diskus. 2002 

articulosus Puplesis & Diskus, 2002 

rotundas Puplesis & Diskus, 2002 

amazonius Puplesis & Diskus, 2002 

insolentis Puplesis & Diskus, 2002 

GLAUCOLEPIS Braun, 1917 

aerifica (Meyrick, 1915) 

argentosa Puplesis & Robinson, 2000 



Fomoria molybditis-group. The group is newly 
designated here for species characterized by the out- 
standingly long transverse bar of the transtilla 
(apomorphy), broad and very long vinculum, and 
bilobed valva (often possessing large spine-like proc- 
esses from the inner side). This distinctive group 
appears to be endemic to the Neotropics and contains 
four species: molybditis (Zeller, 1 877), diskusi Puplesis 
& Robinson, 2000, species 29122, repanda Puplesis & 
Diskus, 2002. The biology of these species remains 

Acalyptris latipennata-group. The group is newly 
designated here for species characterized by a distinctly 
broadened (or moderately broad) forewing with a dark 
oblique fascia, a distinctive paired uncus, and the 
presence of three very large comuti at the apex of the 
aedeagus which tends to be slightly or strongly swollen; 
the valva of species of this group has an inner spine- 
like process (or papilla-like extension). This distinctive 
group appears to be endemic to the Neotropics and 
contains four species: latipenndta (Puplesis & 
Robinson, 2000) (comb.n. - see below), dividua 
Puplesis & Robinson, 2000; ecuadoriana Puplesis & 
Diskus, 2002 and onorei Puplesis & Diskus, 2002; the 
biology of these species is unknown. 

Acalyptris latipennata (Puplesis & Robinson) comb.n. 

Originally this strange-looking species was described 
as a Fomoria (Puplesis & Robinson, 2000). However, 
the later discovery of related species from Amazonian 
rainforest (i.e., ecuadoriana and onorei) indicates, from 
venational and other characters, that this species is 
correctly placed in Acalyptris, to which genus it is here 

Leaf-mine collecting and hostplant data 
in equatorial America 

There has been little elucidation of nepticulid biology 
in the Neotropics (Puplesis & Robinson, 2000). Our 
experience of mine collecting in equatorial America 
suggests that there are great differences between habi- 
tats with respect to diversity and abundance of mines. 
In lowland Amazon rainforest (such as Yasuni, 260 m) 
nepticulid mines are extremely difficult to find, and we 
have never observed mines in abundance although 
adults were readily attracted to light. In disturbed, 
mainly secondary premontane rainforest (Misahualli, 
500 m) mining larvae were much more diverse and 
abundant. In progressing from the foothills of the 
Andes (Tandapi, 1200 m, Banos, 1500-2500) to the 
high Andes (Papallacta, 3500 m, and the slopes of Vol. 
Chimborazo, 4200 m) nepticulid mines became more 
abundant, but the diversity of species dropped signifi- 
cantly above 3000 m. 

Hostplants from only eight plant genera belonging 

to eight families are known for identified species of 
Nepticulidae from the Neotropical region (Table 2). 
Records from Ludwigia and Senecio (Argentina) come 
from a paper by Bourquin (1962) and have not been 
confirmed by other authors. 

We have obtained some additional data from field- 
work in Ecuador, but as rearings produced only single 
female adults or cocoons, these host records remain of 
'unidentified Nepticulidae': 

Rosaceae: Lachemilla: bloch-like mines, cocoons 
cream-white, on southern slopes of Vol. Chimborazo 
(4100-4200 m) (no. 4733-VPU); Acaena: short con- 
torted galleries, on southern slopes of Vol. Chimborazo 
(4100-4200 m) (no. 4734- VPU); unidentified plant 
genus: blotch-like mines, cocoons yellowish, southern 
slopes of Vol. Chimborazo (3500-4000 m) (no. 4737- 

Rubiaceae: Psychotria: sinuous or contorted gallery 
mines, Misahualli, 17 km SE of Tena, Amazon rain- 
forest, 450-500 m (no. 4723, 4743-VPU). 

Fabaceae: Erythrina <?t/«//sTrianaex Micheli. Balu: 
slender sinuous galleries, ochreous cocoons, western 
foothills of Andes, Bucay, ca. 700 m, (Acalyptris sp., 1 
female reared- VPU) (no. 4736-VPU); Inga: long sinu- 
ous leaf-mines (Misahualli 1 7 km SE of Tena, Amazon 
rainforest, 450-500 m) are likely to be also Nepticulidae 
(No 4746- VPU). 

Mines on Bauhinia tarapotensis Benth. (Fabaceae) 
(Amazon rainforest, no. 4630C -VPU) and Cavendishia 
bracteata (Ruiz & Pav. ex J.St. - Hil) Hoerold 
(Ericaceae) (Andes, Banos, ca. 1500 m) (no. 4721 
VPU) cannot be confirmed as of Nepticulidae and may 
have been produced by representatives of other Lepi- 
doptera families. 

Diversity and geography 

Dominance of Acalyptris. One of the most 
unexpected results of investigations of the tropical 
American Nepticulidae was the discovery of a diverse 
fauna of Acalyptris. During the Belize expedition in 
1998, a total of 14 species was found, representing 
48% of Nepticulidae recorded from the area. We 
thought (Puplesis & Robinson, 2000) that the domi- 
nance of Acalyptris might be a regional (Central 
American) or a seasonal phenomenon. But sampling in 
Ecuador (mainly in rainforest) has shown a similar 
pattern exists there and at a different time of the year. A 
total of eight species of Acalyptris was found, repre- 
senting 50% of Nepticulidae recorded from the area. 
Neotropical Acalyptris exhibit a remarkable range of 
morphological structure. 

Table 2. Hostplants of Neotropical Nepticulidae. 


Plant family 

Plant genus 

Nepticulidae reared 


















Enteucha gilvafascia, 
Manoneura basidactyla 
Stigmella gossypii 
Stigmella montanotropica 
Stigmella nubimontana, 
S. rubela 

Enteucha guajavae 
Stigmella guittonae 
Acalyptris species 29140 
Stigmella guittonae 

Davis, 1978 

Forbes & Leonard, 1930 and Davis, 1978 

Cultivated host 
Bourquin. 1962 

Bourquin, 1962 

Endemism of the fauna. The 74 species so far known 
from the Neotropical region all appear to be endemic at 
species or even species-group level. However, only a 
single genus - Manoneura - seems to be endemic to the 
region. Material from the southern part of the continent 
that has been recently studied superficially by the 
authors may contain one or two additional genera. 

Potential diversity. Sampling of nepticulids in the 
Neotropical Region has been sparse, despite the vast 
potential of their habitat, and what we have seen so far 
is probably just a small fraction of the total. The Belize 
sample (January 1 998: 29 species) and Ecuador sample 
(January 2000: 1 7 species) are geographically separated 
by some 2500 km and contain only two species in 
common, Manoneura basidactyla and Ectoedemia 
fuscivittata. The two Amazon rainforest samples 
separated by only 140 km (Yasuni, January 2000: 6 
species; Jatun Sacha, January 2000: 7 species) have 
only a single species in common -Acalyptris insolentis. 
Sampling methods and weather conditions were similar 
at both sites. In the light of this, we would expect 
further collection and study to expand the known 
diversity of Nepticulidae in the Neotropical Region to 
at least 500 species. 

Species numbers by country . The number of species 
known from each neotropical country (Table 1; Fig. 
22) currently indicates only collecting and study 
activity. Only Belize and Ecuador have a species count 
greater than 1 5 . Despite the suggested high diversity of 
Nepticulids in the Neotropics, species counts for 

individual countries are comparable only with the 
most poorly studied countries of other regions. 
They are not comparable with the known species 
diversity of European countries, the product of about 
235 years of investigative history that began with the 
description of Ectoedemia occultella by Linnaeus in 
1767 (Fig. 23). 


Bourquin, F. 1962. Microlepidopteros nuevos con sus hiologias. 

Revista de la sociedad Entomologica Argentina, Buenos Aires, 23: 

3 1^6. 
Davis, D.R. 1978. New leaf-mining moths of the family Nepticulidae 

from Florida. The Florida Entomologist, 61 (4): 209-224. 
Newton, P.J. & Wilkinson, C. 1982. A taxonomic revision of the 

north American species of Stigmella (Lcpidoptera: Nepticulidae). 

Systematic Entomology, 7: 367-463. 
Nieukerken, EJ. van 1986. A provisional phylogenetic checklist of 

the western Palaearclic Nepticulidae, with data on hostplants (Lepi- 

doptera). Entomologica Scandinavica, 17: 1-27. 
Puplesis, R. 1994. The Nepticulidae of Eastern Europe and Asia. 

Leiden. Backhuys Publishers. 291 pp + 840 Figs. 
Puplesis, R. & Robinson G.S. 2000. A review of the Central and 

South American Nepticulidae (Lepidoptera) with special reference 

to Belize. Bulletin of the Natural History Museum. London (Ento- 
mology). 69 (1): 3-114. 
Puplesis, R., Diskus, A. & Robinson G.S. 2002. New Neotropical 

Nepticulidae (Lepidoptera) from the western Amazon rainforest 

and the Andes of Ecuador. Bulletin of the Natural History Museum. 

London (Entomology), 71 (1): 19-58. 
Scoble, M.J. 1983. A revised cladistic classification of the Nepticulidae 

(Lepidoptera) with descriptions of new taxa mainly from South 

Africa. Transvaal Museum Monograph, 2: i-xi, 1-105. 
Zeller, P. C. 1877. Exotische Microlepidoptera. II. Horae Societatis 

Entomologicae Rossicae, 13: 289^193. 






80 km 

i i 

50 miles 

Fig. 1. Map of Ecuador showing collecting localities of 2000 and 2001 fieldwork programmes. 



Figs 2-3. Ecuador collecting sites -Yasuni (240 m): 2, view over canopy (photo: Simon Hill); 3. along theTiputini river. 






'* - •*■■•■■ 


Figs 4-5. Ecuador collecting sites: 4, Misahualli (450 m). premontane tropical forest; 5, Tandapi, western slopes of the 
Andes, montane tropical forest. 




Figs 6-8. Adult Nepticulidae. 6, Manoneura basidactyla, Ecuador; 7, Ectoedemia fuscivittata, Ecuador; 8, Ectoedemia 
fuscivittata, Belize (type locality). 













Figs 9-11. Male genitalia of Manoneura basidactyla, Belize (29120-BMNH): 9, capsule; 10, gnathos, uncus and tegumen; 
1 1, aedeagus. Scale: 0.1 mm. 



Figs 12-15. Genitalia of Manoneura basidactyla, Ecuador: 1 2, male genitalia, capsule (AD0327-VPU); 13, same, gnathos; 
14, same, aedeagus; 15, female genitalia (AD0326-VPU). Scale: 0.1 mm. 




Figs 16-20. Male genitalia of Ectoedemia fuscivittata: 16, holotype, Belize (29107-BMNH), capsule; 17, same, aedeagus; 
18, same, paratype (AD0302-VPU); 19, Ecuador (AD0328-VPU), capsule; 20, same, aedeagus. Scale: 0.1 mm. 



Fig. 21. Distribution map of Nepticulidae species recorded from the Neotropical Region: I . Enteucha cyanochlora; 2, E. 
gilvafascia; 3, E. hilli; 4, E. contracolorea; 5, E. terricula; 6, E. snaddoni; 7. E. guajavae: 8, E. acuta; 9, Manoneura 
basidactyla; 10. M. trinaria; I l,Stigmella andina; 12, S. cuprata; 13, S.johannis; 14, S. ntdis; 15, S. marmorea; 16, S. 
peruanica; 17, S. epicosma; 18, 5. schoorli; 19, S. hamata; 20, 5. imperatoria; 21, 5. olyritis; 22, 5. montanotropica; 23, 5. 
nubimontana; 24, 5. rubeta; 25, 5. eurydesma; 26. S. albilamina; 27, S. fuscilamina; 28. 5. kimae; 29, S. plumosetaeella: 
30, 5. barbata: 31,5. austroamericana; 32, 5. gossypii; 33, 5. pruinosa; 34, 5. ovata; 35. 5. hylomaga; 36, 5. costalimai: 37. 
S. guittonae; 38, Ectoedemia reneella; 39, E. helenella; 40, £. mesoloba; 41, £. species 29105; 42, E.fuscivittata; 43, 
Fomoria tabulosa; 44, F. molybditis; 45, F. diskusi; 46, F species 29122; 47, F repanda; 48,Acalyptris latipennata; 49, A. 
dividua; 50, A. ecuadoriana; 51, A. onorei; 52, A. bovicorneus; 53, A. martinhehngi; 54, A. fortis; 55, A. hispidus; 56, A. 
novenarius; 57, A. lascuevella; 58, A. bifidus; 59, A. trifidus; 60, A. tenuijuxtus; 61 . A. unicornis; 62, A. laxibasis; 63, A. 
bicornutus; 64, A. species 29135; 65, A. platygnathos; 66, A. species 29140; 67. A. basihastatus; 68, A. pseudohastatus; 69, 
A. articulosus; 70, A. rotundus; 7 1 , A. amazonius; 72, A. insolentis; 73, Glaucolepis aerifica; 74, G. argentosa. 





Guyana Colombia 



Fig. 22. Numbers of Nepticulidae recorded from the Neotropics. *The figure for the USA includes only species from 
Arizona and Florida recognized as tropical, and excludes boreal species. 


Great Britain Sweden I Latvia I Ukraine ! Tajikistan South Africa Belize 
Spain Denmark Poland Estonia Turkmenistan Indonesia Ecuador USA 

Fig. 23. Numbers of species of Nepticulidae recorded from various countries. 


Bull. nut. Hist. Mus. Land. (Ent.) 71( 1 ): 77-1 33 Issued 27 June 2002 

A review of the genera associated 
with the tribe Asthenini 
(Lepidoptera: Geometridae: 


Institute of Zoology, Accidentia Sinica, 19 Zhongguancun Road, Beijing 100080, 


Department of Entomology, The Natural History Museum, Cromwell Road, London 
SW7 5BD. U.K. 


Synopsis 78 

Introduction 78 

Comments on the tribal classification 78 

Structures of taxonomic note 79 

Layout 80 

Depositories of material 80 

Acknowledgements 80 

Tribe Asthenini 80 

Checklist of the genera of Asthenini 81 

Genera examined and excluded from Asthenini 8 1 

Key to genera 81 

Asthena 82 

Hydrelia 85 

Agnibesa 89 

Euehoeca 89 

Epicyme 90 

Eschatarchia 91 

'Chalyboclydori flexilinea 91 

Palpoctenidia 92 

Asthenotricha 93 

Venusia 95 

Nomenia 97 

Hastina 98 

Macrohastina 99 

Bihastina 99 

Leucoctenorrhoe 100 

Parasthena 101 

Poecilasthena 101 

Polynesia 104 

Anydrelia 105 

Genera examined and excluded from Asthenini 105 

Minoa 105 

Chalyboclydon 106 

Cleptocosmia 107 

Eois 108 

Pseudopolynesia 108 

© The Natural History Museum, 2002 



Chaetolopha 109 

Trichodezia 109 

References 109 

SYNOPSIS. The Asthenini (Geometridae, Larentiinae) are reviewed at the generic level. Critical 
examination of the composition of the tribe has led to the recognition of 1 8 genera, with the species 
'Chalyboclydon flexilinea Warren unplaced. Species are listed under all of the genera identified 
as Asthenini; their broad distribution is noted; and remarks on peculiarities are noted. Seven other 
genera that have been associated with the tribe are also considered, and reasons for their exclusion 
are given. A lectotype is designated for the non-asthenine species Chalyboclydon marginata 


The primary purpose of this paper is to review the 
genera of the larentiine geometrid tribe Asthenini. 
Most of the genera occur in the Palaearctic and Oriental 
regions, with a few genera in Australia and New Guinea, 
three genera in the Nearctic, one genus in the 
Afrotropics and one in the Neotropics. Representation 
is particularly high in China and the Himalayas. 
Although we are not entirely satisfied with our final 
'definition' of the tribe, we hope, at least, that this 
contribution will provide a material contribution to our 
understanding of part of the asthenine/eupitheciine 
complex. It builds on the extensive study of the Chinese 
species of Larentiinae by Xue & Zhu (1999). The study 
continues those efforts to produce global generic 
reviews of selected groups of Geometridae (e.g., Pitkin, 
1996; Scoble, 1995; Scoble & Kriiger, in press). Given 
the inadequacy of the tribal classification of the sub- 
family, we believe that taxonomic progress is more 
likely to be made at the level of genus and species. For 
examples of recent contributions of this kind see Choi, 
2000; Parra, 1991; Parra & Santos-Salas, 1991; 
Schmidt, 2001; Xue & Zhu, 1999). 

We have two specific aims. First, given the lack of an 
explicit definition of the Asthenini, the tribe is examined 
critically for coherence, so we use this paper as a 
means of exploring taxonomic problems. Our concept 
of the tribe is presented through the description, 
diagnosis and generic checklist, which follow a section 
in which significant taxonomic characters are reviewed. 
Second, despite the shortcomings of the tribal 
definition, we provide a treatment of the genera and 
state our reasons for their inclusion in the tribe. Certain 
genera that have been associated with the Asthenini in 
the past, or that we ourselves consider close to the 
tribe, are treated at the end of this work with an 
explanation as to why they have been excluded from 
the Asthenini. 

Comments on the tribal classification 

In reviewing the larentiine tribe Asthenini, we were 
faced with a question common to virtually all global 
taxonomic treatments: how inclusive should we be in 
associating genera with a higher taxon originally 
defined from relatively few European species? Our 
approach has been to compare as many potential 
asthenine taxa as possible against the existing literature 
and discuss their inclusion in, or their exclusion from, 
the tribe. 

A difficulty in deciding which genera to include was 
that the tribal classification of Larentiinae as a whole 
remains significantly unresolved. It has developed 
largely from a series of regional studies. The Asthenini 
were established (as subfamily Astheninae) by Warren 
(1893: 362). Besides associating Asthena, Hydrelia 
and Chalyboclydon he included a part of the 
Trichopterygini, but he gave no definition or description 
of the tribe. 

Although L.B. Prout, in his studies of larentiine 
genera, also never actually described the tribe, many of 
the genera we associate with the Asthenini today were 
treated in reasonably close proximity (e.g., in the 
sections on Larentiinae in Prout, 1912-1916). The 
group (as 'Astheninae') was more effectively founded 
by Pierce (1914: 38), whose diagnosis rested on 
characters of the genitalia, but included just the genera 
Minoa and Asthena, for the study was restricted to the 
British fauna alone. Pierce's diagnosis was based on a 
reduced uncus, entirely attached to the anal tube, the 
presence of an extended valval sacculus in the male, 
and a long evenly spined signum on the corpus bursae 
of the female. Pierce also noted the presence of labides 
(arm-like sclerites of the diaphragma) in each species 
included in the tribe, but did not include these structures 
in his tribal diagnosis, presumably because they occur 
elsewhere in Larentiinae, notably in Eupitheciini. 

McGuffin (1958), still in a pre-cladistic age, and in 
a study restricted to larentiine larvae, published a 



figure in which Asthenini were represented as the most 
basal group of Larentiinae from which four other tribes 
arose directly. In cladistic terminology, the group would 
be viewed as paraphyletic and, therefore, unsatisfac- 
tory. An examination of McGuffin's text, however, 
suggests that he perceived the group as more 
phylogenetically coherent. McGuffin restricted his 
study to Larentiinae of North America and included 
just the two genera Hydrelia and Venusia. He suggested 
that larval morphology demonstrated a close relation- 
ship between the two genera. The spinneret was found 
to be much longer than the labial palpi, and the thoracic 
claw was described as being almost straight, with the 
angle of the notch being acute. How far these observ- 
ations pertain to the Asthenini more widely, and whether 
the characters are apomorphic, remain unknown. 
Larvae of most Asthenini are unknown, and our work 
has been based, inevitably, on adult morphology. 

McGuffin's classification was preceded by that of 
Forbes (1948), which was also restricted to North 
American Asthenini. Forbes included three genera, 
Hydrelia, Venusia and Trichodezia, but apart from 
some comments in a 'table of tribes', he did not give a 
convincing definition of the tribe. He seems to have 
based his concept of the group largely on the existence 
of extended chaetosemata. We have excluded 
Trichodezia from Asthenini, for it has a well-developed 

A list of family-group names of Larentiinae was 
compiled by Holloway ( 1 997 ) in his work on the moths 
of Borneo. Holloway did not distinguish the Asthenini 
from the Eupitheciini because some of the genera that 
had been added to the group by McQuillan & Edwards 
(in Nielsen, Edwards & Rangsi, 1 996) lack an extended 
sacculus, a character used to define the tribe by Pierce 
(1914), and because he considered the distinction 
between Asthenini and Eupitheciini to be weakly 

Chinese species of Larentiinae were treated by Xue 
& Zhu (1999) in their extensive survey of the sub- 
family. Many of the genera included in the present 
paper were discussed there, but tribal definitions were 
not provided. 

Structures of taxonomic note 

Uncus, tegumen and anal tube. Typically in 
Lepidoptera, the anal tube is attached to the tegumen 
posteriorly and, where an uncus is present, the anal 
tube diverges at the point at which it articulates with 
the tegumen. In his description of the Asthenini (as 
Astheninae), Pierce (1914: 38) stated that 'the uncus is 
weak and entirely attached to the anal tube which bears 
a thickened subscaphium'. In most species, the uncus 
(a sclerite articulated with the posterior end of the 
tegumen) appears to be absent. Occasionally (as in 
Eschatarchia) a triangular vestige is apparent. In the 

light of this observation, it is difficult to understand as 
a general observation Pierce's statement that the uncus 
is attached to the anal tube in Asthenini. However, 
where a vestige is present, it is indeed fused entirely 
with the dorsal surface of the anal tube. 

Sclerotizations associated with the diaphragma and 
anellus. Although the sclerotizations of this region 
are complex within the Larentiinae and much used in 
the taxonomy of the subfamily, little has been written 
on the homologies of the various components. There is 
a real need for a comparative study of the area across 
the group. The important sclerotizations in the 
Asthenini and Eupitheciini are the labides, the 
transtillae and the juxta. 

Each transtilla extends from the base of each valva 
dorsally. In most Asthenini (and, indeed, across the 
Lepidoptera) these paired structures meet medially 
and represent the dorsal-most sclerotization of the 
diaphragm. The transtillae are often not very conspicu- 
ous, particularly where the genitalia are mounted 

The labides were said by Pierce (1914) to spring 
from the points at which the transtillae unite with the 
costae of the valvae. This is a somewhat narrow view, 
since there are a number of rod-like structures in 
Larentiinae that may or may not be homologous with 
these structures in Asthenini - in, for example, 
Eupitheciini. Of particular interest in this paper is the 
comparison between the Eupitheciini and the Asthenini. 
In Asthenini, the labides usually arise from the base of 
the valvae, as noted by Pierce, and extend variously 
(see generic treatments, below). In Eupithecia, it does 
not seem to be have been stated that labides homolo- 
gous to those in Asthenini actually exist. However, 
observations on the genus Poecilasthena have shed 
some light on this matter (see below). 

In Eupithecia the juxta, possibly with other 
sclerotizations of the diaphragm, is distinctive being 
shaped like an hourglass (see, e.g., illustrations in 
Holloway, 1997). The anterior end of each member of 
a pair of ventral arms lateral to the juxta curve inwards 
to the medial constriction of the juxta. Structures that 
appear to be articulated with the posterior ends of the 
arms meet medially to form what are possible homo- 
logues of the asthenine labides. 

In Poecilasthena although the juxta is flask-shaped, 
not hourglass-shaped, there occurs what appears to be 
the homologue of the ventral arms in Eupithecia. Each 
arm continues anteriorly into a somewhat expanded 
and free membranous structure, which is not united 
medially with its opposite member. We consider that 
the arms and their membranous expansions are labides, 
even though they are not united with the base of the 
valvae as they are in typical Asthenini. Support for this 
view comes from observations on P. paucilinea, in 
which what we take to be the ventral arms do extend 


from the bases of the valvae, and are more strongly 
sclerotized than normal and less arm-like. Females of 
both Poecilasthena and Eupithecia bear a small 
colliculum within a narrow ductus bursae. Other 
features of Poecilasthena, are, however, much closer 
to the Asthenini condition, notably the shape of the 
juxta and the signum. Under this interpretation, the 
ventral arms in Eupithecia are also part of the labides. 
Our study of Poecilasthena gives further support to 
the view that the taxonomic association of the Asthenini 
and Eupitheciini is close. Indeed, like Holloway ( 1 997), 
who included Parasthena, Poecilasthena, Polynesia, 
Eois, and Pseudopolynesia in a broad definition of 
Eupitheciini, we harbour some doubts that the tribes 
are distinct. 

Venation (Figs 189-192). In most Asthenini the 
forewing areole is single, but examples of a double 
areole occur (as in most Venusia, in Bihastina, and in 
Poecilasthena), and in 'Chalyboclydon flexilinea and 
Palpoctenidia the areole is absent. Differences between 
genera also occur in the point at which veins R ( and R. 
diverge from the common stem in the forewing. In some 
genera, R ( diverges before (proximal to) the divergence 
point of R 5 , whereas in others the opposite condition is 
encountered. In the hindwing, the discocellulars are 
either markedly angled (biangulate condition) or not so 
modified. The position of vein M, in relation to veins M 
and M , and whether or not M is united at its base 
(stalked) with CuA : are other venational features found 
to have some taxonomic value. 

Corpus bursae. The typical asthenine signum is 
composed of a dense accumulation of denticles or 
spines radiating, on both sides, from a central line or 
ridge formed from the bases of the denticles or spines. 
An example of a signum showing a combination of 
denticles, radiating spines and a central ridge is 
illustrated in Fig. 295 (Poecilasthena dimorpha). The 
signum is usually elongated, but sometimes round or 
elliptical. In some Asthenini it takes the form of a 
naiTow band. The signum in Asthenini is distinctive, 
providing, probably, the best defining character for the 
group. In Hydrelia, besides the main signum, a smaller 
second signum occurs, which is also formed from 
denticles arranged in the same way. In Palpoctenidia 
the spines are stouter than usual, but the radiating 
arrangement is the same. In the non-asthenine genus 
Pseudostegania Butler, a signum composed of 
numerous denticles occurs. However, the arrangement 
of these differs from the condition in Asthenini for the 
denticles do not radiate from a central line. The signa 
in Sterrhochaeta Prout also differ. (The type species of 
this genus, S.fulgurata (Warren) is illustrated in Fig. 

In a number of species of Asthenini, the corpus 
bursae is partly or wholly covered with denticles. 
Often these denticles are minute, but in some species 
they are prominent (e.g., as in Fig. 277) with strength- 
ening ridges. These denticles are frequently 
encountered in Eupithecia, but they occur in other 
genera of Larentiinae such as Horisme (see, for 
example, in Holloway. 1997), in some Sterrhinae and 
outside the Geometridae. In the Asthenini, these 
denticles do not occur across individual genera, but are 
present sometimes in just one species. The taxonomic 
distribution of these denticles, therefore, renders it 
difficult to draw wide phylogenetic conclusions from 
their presence. Nevertheless, when other characters 
shared by Eupithecia and Asthenini are taken into 
account, the presence of these denticles lends further 
support for a close association of the taxa. 


A list of species included in each genus of Asthenini is 
presented with the name of the original genus, if differ- 
ing from that current, provided in brackets. We have 
also listed species belonging to some of the non- 
asthenine genera that are discussed at the end of this 
paper. References to species listed have not been cited 
as they are to be found in Scoble (1999), but they are 
given for type species, which are detailed under the 
description for each genus. 

Depositories of Material 

ANIC The Australian National Collection, Canberra. 

BMNH The Natural History Museum. London, UK 
IZAS Institute of Zoology, Academia Sinica. Beijing. 


Acknowledgements. The illustrations were compiled 
by Shayleen James and Maia Vaswani (BMNH). to whom we 
are extremely grateful for their competence, patience and 
willing support. Shayleen James also prepared digital images 
of the whole moths and arranged these and those of the 
genitalia, which were taken by XD, into plates. For valued 
discussion on this work we thank our colleagues from the 
BMNH. or associated with this institution, namely Jeremy 
Holloway. Ian Kitching, Linda Pitkin and Tony Galsworthy. 
We are particularly grateful to Jeremy Holloway for his 
comments on a draft of the text. Our thanks are also due to 
Axel Hausmann (Zoologische Staatssammlung, Munich) and 
Olga Schmidt (Staatliches Museum fiir Naturkunde, Stutt- 
gart) for further discussion on aspects of the work. 


Astheninae Warren, 1893: 362. 

MOTH(Figs 1-177). Generally fairly small compared 
with other Larentiinae. Body typically slender. Head. 



Frons often broad and protuberant, but sometimes 
neither broad nor protuberant. Labial palpi narrow and 
short, generally not pointed or only weakly pointed. 
Antenna of male ciliate or smooth, sometimes 
bipectinate, seldom unipectinate. Legs simple: tibial 
spurs 0-2—4. 

Wings generally pale with dark waved transverse 
lines running from costa of forewing to hind margin of 
hindwing. Venation as in Figs 189-192. Forewing: 
usually with areole single, sometimes double, occasion- 
ally absent; if areole is absent, vein R never connected 
to Sc, in contrast with some Eupitheciini. Hindwing: 
hind margin generally narrow; Sc+R : diverging from 
common stem 3/4 of way along cell; Rs sharing com- 
mon stem with M : ; 3 A absent. 

MALE GENITALIA (Figs 193-220; 226-253). Uncus 
absent or strongly reduced. Anal tube united with 
tegumen (and uncus vestige, if present); subscaphium 
often present. Vinculum: extended into broad or nar- 
row saccus. Labides fairly membranous, apex setose. 
Juxta often broad at base, tapering - sometimes to a 
slender process posteriorly. Valva: broad, setose, with 
setae pointing towards base of costa; sacculus usually 
bearing sclerotized extension, extension sometimes 
not developed. Anellus: labides present, varying from 
narrow to broad, or absent; juxta broad or narrow. 
Aedeagus: narrow; vesica with various sclerotizations. 

FEMALE GENITALIA(Figs 268-292). Ductus bursae 
of constant width, often sclerotized extensively; broader 
than in Eois. Corpus bursae seldom densely denticu- 
late, unlike the condition in Eupitheciini and 
Trichopterigini, if denticles occur, they do not 
completely cover corpus bursae; signum distinctive, 
composed of denticles or spines radiating from central 
line or ridge formed by their bases; usually somewhat 
elongated, sometimes in form of narrow band, some- 
times almost round; second smaller signum, of same 
basic composition, sometimes present. 

DIAGNOSIS. Asthenines are best distinguished from 
other Larentiinae by the presence of the distinctive 
signum. Asthenini share some characters with the tribe 
Eupitheciini. The combination of extremely narrow 
and short labial palpi, a generally broad and protuber- 
ant frons and relatively broad wings are typical external 
features of Asthenini. The best diagnostic combination 
of characters of the genitalia includes the reduced or 
lost uncus; labides narrow, rather than lobe- or spoon- 
shaped, and seldom united; and the corpus bursae with 
a signum (or signa) composed of radiating denticles or 

Distribution. Asthenini are represented in all the 
major biogeographical regions. The tribe is at its most 
I diverse in east Asia, but is poorly represented in the 
Afrotropics and Neotropics. 

Taxa INCLUDED. Eighteen genera are included and 
the species 'Chalyboclydon' flexilinea Warren is 
unplaced. Two hundred and twenty-seven species group 
taxa (species and subspecies) are identified as belong- 
ing to the tribe. 

MATERIAL EXAMINED. About 2900 specimens have 
been studied: 2558 from the BMNH, and 350 from 
IZAS. Approximately 400 genitalia slides from the 
BMNH collection were examined. 

Checklist of the genera of Asthenini 

Asthena Hiibner, [1825] 1816 
HydreliaUiibaer[l&25] 1816 
Agnibesa Moore. 1 888 
Euchoeca Hiibner, [1823] 1816 
Epicyme Meyrick. 1 885 
Eschatarchia Warren, 1894 
'Chalyboclydon' flexilinea Warren, 1 898 
Palpoctenidia Prout, 1930 
Asthenotricha Warren. I 899 
Venusia Curtis, 1X39 
Nomenia Pearsall, 1905 gen. rev. 
Hastina Moore. 1888 
Macrohastina Inoue, 1982 
Bihastina Prout, 1916 
Leucoctenorrhoe Warren. 1904 
Parasthena Warren. 1902 
Poecilasthena Warren. 1 894 
Polynesia Swinhoe, 1892 
Anydrelia Prout, L938 

Genera examined and excluded from 

M inoa Treitschke, 1825 
Chalyboclydon Warren. 1893 
Cleptocosmia Warren, 1896 
Eois Hiibner, 1818 
Pseudopolynesia Holloway. 1997 
Chaetolopha Warren. 1 899 
Trichodezia Warren. 1895 

Key to Genera 

1. Forewing usually with R, diverging from stem distal to 
divergence point of R : if R does not diverge in this way, 
then either cell of hindwing is no longer than 1/3 of length 
of hindwing, or discocellulars of forewing are strongly 
biangulate 2 

Forewing with R f diverging from stem proximal to diver- 
gence point of R5. or united only briefly, or diverging at 
same point: hindwing cell extending almost to half length 
of hindwing or beyond; discocellulars of forewing never 
biangulate 12 

2. Cell of hindwing not longer than 1/3 length of hindwing 


Cell of hindwing close to or exceeding half length of 
hindwing 4 



3. Forewing with areole absent . 


Forewing with areole single Polynesia 

4. Discocellulars strongly biangulate on forewing; male 

antenna unipectinate, with short pectinations 


Discocellulars not biangulate on forewing. male antenna 
never unipectinate 5 

5. Antenna in both sexes bipectinate Leucoctenorrhoe 

Antenna not bipectinate in either sex 6 

6. Discocellulars of hindwing strongly biangulate; male 
hindwing with hind margin expanded into very large flap 
which is folded beneath wing and almost reaches to 
middle of cell Anydrelia 

Discocellulars of hindwing not biangulate; male hindwing 
without this modified flap 7 

7. Termen of hindwing with sharp angle, turned upwards at 
M ; both wings with fuscous line near the distal margins, 
that of forewing bent strongly, touching the distal margin 
medially and enclosing pale marginal patches on upper 
and lower surfaces Eschatarchia 

Termen of hindwing not angled as above; wings lacking 
fuscous line near distal margins 8 

8. Termen of forewing dentate, deeply excavated between 
R,andM, 9 

Termen of forewing gently curved, lacking tooth or deep 
excavation 10 

9. Frons prominent, broad and rounded; wings dark or pale 
brown with yellowish lines; vein M, never stalked with 
CuA! Hastina 

Frons not prominent, nor broad nor rounded: apical area 
of forewing and distal part of hindwing partly pure white, 
proximal half of hindwing pale yellowish; vein M some- 
times stalked with CuA : Macrohastina 

10. Wings pale greyish to white; hindwing with termen 
excavated between M and M , points at M and M sharp 


Wings yellowish brown: hindwing not excavated as 
described above and lacking point at M ; . termen angled at 
M, but blunt, not sharp Euchoeca 

11. Small (length of forewing not longerthan 10mm); wings 
pale greyish, lacking visible white ground colour; male 
forewing with small anal lobe, male hindwing with 
posterior margin folded under wing Parasthena 

Large (length of forewing not less than 13 mm); white 
ground colour of wings always visible; male wings lack- 
ing such modifications Agnibesa 

12. Forewing lacking areole; male antenna bipectinate 


Forewing with 1 or 2 areoles; male antenna not bipectinate 

13. Discocellulars of hindwing biangulate 14 

Discocellulars of hindwing not biangulate 15 

14. Forewing with areole single Venusia 

Forewing with areole double 

Poecilasthena (main group and anthodes group) 

15. Termen of hindwing deeply dentate Bihastina 

Termen of hindwing not dentate, smooth or slightly 
waved, or angled at middle 16 

16. Forewing with areole usually double, occasionally single; 
if areole single, wings white; if areole double, forewing 
with antemedian, median and double postmedian lines 
dark brown and running straight below costa: vesica with 
bundle of cornuti on vesica 17 

Forewing with areole single, markings not as above; 
vesica without cornuti 18 

17. Ground colour of wings pale grey; termen of hindwing 
weakly dentate at M, and M,, excavated between teeth; 
male genitalia with coremata; vesica lacking cornuti; 
proximal edge of sternum A8 of male modified to form 
'W shape Poecilasthena (papuensis group) 

Ground colour of wings white, termen of hindwing weakly 
angled medially, seldom dentate and excavated; male 
genitalia lacking coremata: vesica with cornuti present; 
sternum A8 unmodified Asthena 

1 8. Costa of forewing strongly expanded proximally. or male 
hindwing with proximal half of costa strongly broad- 
ened: hair tuft present on upper surface of costa close to 
base; anastomosis of Sc+F^ with Rs distinctly less than 3/ 
4 length of hindwing; uncus/tegumen triangular and 
pointed; labides finger-like, extending to half length of 
tegumen; anal papillae nearly smooth, distal half of ductus 
bursae 'Y' shaped 19 

Costa of forewing straight or only very weakly broadened 
near base, male hindwing lacking hair tuft; anastomosis 
of Sc+R, with Rs reaching beyond 3/4 length of hindwing; 
genitalia variable, but not as described above 20 

19. Male hindwing with hair tuft on upper surface of costa 
close to base; proximal half of hindwing costa strongly 
broadened Asrhenotricha {dentatissima group) 

Male hindwing lacking hair tuft; hindwing costa not 
broadened Asthenotricha (argyridia group) 

20. Frons not broad and protuberant; valva with apex fringed 
with hair-like scales, which are long and expanded at tips; 
posterior half of ductus bursae membranous, signum 
elliptical, with long spines radiating peripherally from 
denticulate core Epicyme 

Frons generally broad and distinctly protuberant: if not 
like this, then genitalia of both sexes different from above 

Asthena Hiibner, [1825] 1816 

(Figs 1-15; 189; 193-195; 226-228; 268,269.) 

Asthena Hiibner, [1825] 1816: 310. Type species: 
Geometra candidata [Denis & Schiffermuller], 
1775: 1 10 (a junior synonym of Asthena albulata 



Roessleria Breyer, 1869: xix. Type species: Geometra 
candidata [Denis & Schiffermiiller], 1775: 1 10. 

Moth (1-15). Head. Frons neither broad nor 
protuberant. Labial palpi very slender and short, gen- 
erally not extending beyond front of head. Antenna of 
male serrate, with hair tuft in type species and 
nymphaeata, or smooth. Wings. White or off-white in 
most species with pale brown, weak fasciae; appear- 
ance pale brown in e.g., plenaria; mottled brown in 
albosignata; fasciae bold in opedogramma and 
tchratchraia. Venation (Fig. 189) with cell on both 
wings extending slightly less than half length of wing; 
forewing with areole usually double, sometimes single; 
vein R ( arising proximal to apex of second areole; R 5 
arising from apex of second areole or united for short 
distance with R ; hindwing with discocellulars slightly 
curved, not biangulate. distal margin slightly angled at 

MALE GENITALIA (Figs 193-195; 226-228). Labides 
in form of well-developed, straight arms, sometimes 
reduced to very short processes, sometimes absent. 
Juxta broad, extending anteriorly into a narrow process 
so that whole appears flask-shaped. Valva: sacculus 
usually with distinctive double projection of a longer, 
narrow spine-like process and a shorter, broader process 
with short hairs at apex, notably complex in amurensis; 
costa margin weakly convex. Aedeagus: vesica with 
spine-like cornuti. Abdomen: sternum A8 unmodified. 

Female genitalia (Figs 268,269). Ductus bursae 
sclerotized throughout length except for very short 
membranous section. Corpus bursae: signum generally 
short and broad, composed of radiating denticles; 
spinose patch often occurring in addition to discrete 
signum; surface of corpus often partly covered with 
minute denticles. 

Diagnosis. Most species of Asthena may be distin- 
guished from species in other asthenine genera by the 
white ground colour of the wings, but this feature is not 
universal within the genus. Both anseraria and lassa 
resemble superficially certain Scopula species 
(Sterrhinae). Distinguishing characters include the fact 
that the frons is neither broadened nor protuberant, and 
the unstalked condition of vein R ( of the forewing. The 
i most distinctive character is the complex form of the 
sacculus with its two projections. The presence of 
cornuti on the vesica of Asthena distinguishes it from 
Hydrelia where these structures are absent. 


Across the Palaearctic region and in 

Species included. Twenty-two described. There is 
an undescribed species from Burma represented by a 
single, worn male specimen in the BMNH. Genitalia 
examined: A. albosignata (male, female), albulata 
(male, female), amurensis (male), anseraria anseraria 

(male, female), anseraria corculina (male, female), 
hamadryas (male), lassa (male), nymphaeata (male, 
female), opedogramma (male, female), plenaria 
(male), sachaliensis (female), melanosticta (male, 
female), octomacularia (female), tchratchraria 
(female), undulata (male, female). 

Asthena albidaria (Leech, 1897) 

Asthena albosignata (Moore, 1888) (Idaea?) 

India, China. Kashmir. 

Asthena albulata (Hufnagel, 1767) 

Geometra candidata [Denis & Schiffermiiller], 1775 

Widespread in the Palaearctic. 

Asthena amurensis (Staudinger, 1897) 

Cidaria candidata amurensis Staudinger, 1897. 
Asthena hamadryas Inoue, 1976. Syn. n. 

Russia, Japan, Korea. 

Asthena anseraria anseraria (Herrich- 
Schaffer, 1855) (Arrhostis?) 

Cidaria solclaria Turati, 1879. 
Widespread in the Palaearctic. 

Asthena anseraria corculina Butler, 1878 

Japan, China. 

Asthena lactularia (Herrich-Schaffer, 1855) 


? albeolata Rambur, 1 866 

Asthena nymphuluta Guenee, [1858] 

France, Spain. 

Asthena lassa Prout, 1926 


Asthena livida (Warren, 1896) comb. n. 



Remarks. This species differs from others in 
Asthena, and its position in the genus is uncertain. The 
moth is uniformly dark brown rather than white or 
otherwise pale. Furthermore, the cell on both fore- and 
hindwing is very short - on the hindwing it is only 1/3 
the length of the wing. The free section of vein R t of the 
forewing is unusually long. The male genitalia are 



unusual in having an elongated tegumen and narrow, 
but well developed, curved labides. Both these 
characters are reminiscent of the condition in Poly- 
nesia, but the wing pattern is completely different and 
the saccus is not truncated as in that genus. 

Asthena melanosticta Wehrli, 1924 


Asthena nymphaeata (Staudinger, 1897) 

Acidalia ainoica Matsumura, 1927. 

Russia, Japan, Korea, China. 

Asthena ochrifasciaria Leech, 1897 


Asthena octomacularia (Leech, 1897) 

China, Japan. 

Asthena opedogramma (Prout, 1926) comb. 

n. (Hydrelia) 

Burma, China. 

Remarks. This species and tchratchraria Oberthur 
were treated as members of Hydrelia by Prout (1926, 
1934-39) probably because Prout thought that the 
areole in the forewing was always single in that genus, 
whereas in Asthena it is double. Having checked all 
specimens of both species in the BMNH and IZAS, we 
discovered that the number of areoles varies. In some 
specimens just a single areole is. indeed, present. But 
in others, the areole is double on both the right and the 
left forewing, and in yet others it is double on just the 
right or the left fore wing. When the areole is double, 
the first (more proximal) areole is very small. The two 
species share most characters of typical Asthena other 
than the presence, sometimes, of a single areole. The 
frons is only weakly protuberant, cornuti are present, 
and the signum is short and broad, as in many Asthena 
species. Unique to the species is the presence of well- 
sclerotized spines on the lateral parts of the juxta, 
labides and terminal half of the sacculus. 

Asthena plenaria (Leech, 1897) (Hydrelia) 

Asthena sachaliensis (Matsumura, 1925) 

Japan, Russia. 

Asthena tchratchraria (Oberthur, 1893) 


Burma, China. 

(See Remarks under opedogramma Prout.) 

Asthena undulata (Wileman, 1915) 



The albifera group 

The following four species have a single areole, but the 
genitalia are very similar to Asthena notably in the 
shapes of the sacculus, juxta, and vinculum. Additional 
spining occurs on the corpus bursae in addition to the 
denticulate signum. 

The albifera group differs from Hydrelia in having 
a pure white ground colour, a narrow and non- 
protuberant frons, and a bundle of cornuti on the vesica 
of the aedeagus. All of these characters correspond to 
those in Asthena. 

Genitalia examined: albifera (male, female), 
chionata (male), percandidata (male, female). 

Asthena albifera (Walker, 1866) (Acidalia?) 

Acidalia albogilvaria Morrison, 1874 
Corycia triseriata Packard, 1 874 

North America. 

Asthena brunneifasciata (Packard, 1876) 


Asthena chionata (Lederer, 1870) (Cidaria) 

Cidaria quadripunctata Biernert, [1871] 


Asthena percandidata (Christoph, 1893) 


Cidaria anseraria candidissima Staudinger, 1897 
Transcaucasus, Central Asia. 

Species excluded 

The following species, which have been associated 
with Asthena in the past, belong neither to that genus, 
nor to the tribe Asthenini. Genitalia examined: 
argentipuncta (male, female), argyrorrhytes (male, 
female), aurantiaca (male), eurychora (female), 
straminearia (male, female), yargongaria (male). 

'Asthena' argentipuncta Warren, 1906 


Papua New Guinea. 

'Asthena' argyrorrhytes Prout, 1916 


Irian Jaya. 

'Asthena' aurantiaca Prout, 1926 (Asthena) 
Irian Jaya. 

'Asthena' distinctaria (Leech, 1897) 



'Asthena' eurychora Prout, 1928 (Asthena) 
Western Samoa. 

'Asthena' straminearia (Leech, 1897) 



'Asthena' subditaria Warren, 1906 (Asthena) 
Papua New Guinea. 

'Asthena' yargongaria Oberthiir, 1916 



Hydrelia Hiibner, [1825] 1816 

(Figs 16-68; 196,197; 229,230; 270.) 

Hydrelia Hiibner, [1825] 1816: 322. Type species: 
Geometra sylvata [Denis & Schiffermtiller], 1775: 

Autallacta Warren, 1 893: 365. Type species: Timandra 
subobliquaria Moore, 1868: 644. 

MOTH (Figs 16-68). Head. Frons broad and 
prominent. Labial palpi very slender and short, 
generally not extending beyond front of head. Antenna 
weakly serrate, ciliated in male, simple in female. 
Wings. Rather narrower than in Asthena and Venusia. 
Colour variable: most species grey-brown with weak 
transverse band and lines; some species ochreous; 
strong transverse line or lines in a few species; some 
species with yellow ground colour; a few species with 
wings dark grey-brown with contrasting white mark- 
ings. Forewing with single areole, varying in size 
among species; vein R ( diverging from common stem 
before (proximal to) point of divergence of R.. 
Hindwing: termen sometimes angled medially; 
discocellulars not biangulate; vein M usually not 
stalked with CuA r but stalked on both wings in a few 

Male genitalia (Figs 196,197, 229,230). Labides 
in form of a pair of long, curved, spine-like processes 
in type species and relatives; processes smaller and 
membranous in other species or united with opposite 
member. Juxta variable; narrow and extended, plate- 
like or reduced to small sclerite between the bases of 
the valvae. Valva usually narrows to apex; sacculus 
typically extended into a thumb-like projection, some- 


times sacculus not extended. Aedeagus: vesica lacking 
cornuti. Sternum A8 unmodified. 

Female genitalia (Fig. 270). Anal papillae 
extended to form more attenuated ovipositor in type 
species and relatives compared with other species, 
where ovipositor is much flatter. Ductus bursae: antrum 
sclerotized throughout apart from a narrow 
membranous break. Corpus bursae often with numer- 
ous minute denticles; signum typically elongated, 
composed of radiating denticles; many species with a 
small additional signum, similarly composed, at 
posterior end of corpus; surface of corpus partly or 
wholly covered with minute denticles. 

DIAGNOSIS. Close to Asthena, Venusia and Agnibesa. 
Hydrelia differs from Asthena in having a generally 
broad and strongly prominent frons, in the ground 
colour of the wings seldom being pure white, in having 
a single areole on the forewing, in lacking cornuti on 
the aedeagus, and in having a signum typically 
elongated and often band-like. Hydrelia lacks 
biangulate discocellulars on the hindwing and in this 
way differs from Venusia. It may be distinguished from 
Agnibesa by several features (see Diagnosis of 

Distribution. Palaearctic, Nearctic and Oriental 

Remarks. The integrity of Hydrelia as a mono- 
phyletic entity remains in doubt, there being no very 
convincing apomorphy. Wing shape and pattern assist 
recognition of the genus as it stands now. The male 
genitalia of the type species H. sylvata (Fig. 197), and 
its relatives differ from those of other species (e.g., as 
in Fig. 196) and it may eventually be necessary to 
restrict the genus Hydrelia to this group. We do not 
make such a restriction here since Hydrelia is suffi- 
ciently coherent and well known to make it 
counterproductive to split it until a full species-level 
revisionary study has been undertaken. 

Preliminary study has identified several species 
groups, although we do not treat them as formal taxa at 
this stage, and nor are the suggested divisions compre- 

The sylvata group. The type species of Hydrelia, H. 
sylvata (D. & S.), which is similar to the Nearctic 
species H. lucata (Guenee), bears a pair of long, narrow 
and curved labides. The juxta of these species unites, 
anteriorly, with a long and narrow dorsal sclerite of the 
diaphragma. The labides are broader and straight in 
flammeolaria (Hufnagel). That sylvata and 
flammeolaria, both European species, are closely 
related is evident from the relatively attenuated ovi- 
positor in each and in the similar shape of the valva. 
The larval foodplant range overlaps, in that both species 
occur on Alnus (Betulaceae). The Nearctic species 
condensata, inornata and lucata also belong to this 


group. Oriental species that appear to belong with 
sylvata and its relatives include aurantiaca, rufigrisea, 
sericea, nepalensis, rubrilinea, llineata, llaetivirga, 
rhodoptera, marginepunctata, binotata, rubricosta, 
sanguiniplaga, nisaria. The following species, which 
exhibit similarities in wing shape and markings may 
also be associated, or, may form a separate group: 
sericea, rubrilinea, lineata, laetivirga. 

The aggerata group. The largest of these species 
groups, which is Oriental in distribution (mainly Chi- 
nese), is a monophyletic assemblage including aggerata 
and its relatives and is defined by several characters. 
The labides seem to be absent and should not be 
confused with a pair of setose, membranous projec- 
tions from the diaphragma. An additional feature, giving 
some further possible support to the grouping is that 
the valva is broad with the sacculus projected in such a 
way as to form a wide, c. 90" angle, with the costa. 
Unlike the condition in the sylvata group, the oviposi- 
tor is short and not pointed. Species belonging here 
include: aggerata, aurantiaca, bella, bicolorata, 
conspicuaria, crocearia, ornata, pavonica, rubrivena 
and subobliquaria. 

The ungularia group (Oriental) has a rounded valva 
that lacks a projection of the sacculus. The dorsal 
sclerite of the diaphragma is absent. Species identi- 
fied: latsaria, microptera, subtestacea and ungularia. 

The valva of H. impleta Prout differs in shape. 

In subobliquaria (Moore) from Bengal, the valva 
resembles more closely that of the sylvata group than 
that of aggerata and its relatives, but the labides, 
although distinct, are shorter and membranous. 

Species included. 6 1 species. Genitalia examined: 
aggerata (male), aurantiaca (male, female), bella 
(male, female), bicolorata (male, female), binotata 
(male, female), condensata (female), conspicuaria 
(male, female), controversa (male, female), crocearia 
(male, female), elegans (female), enisaria (male), 
fuscocastanea (male, female), impleta (male), inornata 
(female), laetivirga (male), latsaria (male), lineata 
(male, female), lucata (female), marginepunctata 
(male, female), microptera (male, female), nepalensis 
(male, female), ornata (male), pavonica (male), 
rubricosta (male), rubrilinea (male, female), rubrivena 
(male, female), rufinota (female), sericea sericea (male, 
female), subobliquaria (male, female), rhodoptera 
(male, female), speciosa (male), sanguiniplaga (male), 
subcingulata (male), subtestacea (male), sylvata 
(male), undularia (male). 

Hydrelia aggerata Prout, 1938 


Hydrelia arizana (Wileman, 1911) (Acidalia) 

Hydrelia aurantiaca Hampson, 1903 

China, Nepal. 

Hydrelia bella (Wileman, 1916) sp. rev. 



Remarks. The species was listed as a synonym of 
bicolorata (Moore) in Scoble (1999). but the genitalia 
are distinct and it is treated here as a separate species. 
Male with valva much narrower than in bicolorata 
(Moore), shape intermediate between that species and 
aggerata Prout from W. China. In the female, the 
signum in bella is possibly larger than that in bicolorata. 
Wileman (1916: 97) cited the type material as: 'A 
male specimen from Arizan (7300 ft.), September 
27th, 1906, a female specimen from Arizan, August, 
1908; and another from Rantaizan, May, 1909 (7500).' 
The 'male' syntype is, in fact, a female, and there is no 
specimen of the male sex in the type series. 

Hydrelia bicauliata Prout, 1914 

Japan, China. 

Hydrelia bicolorata (Moore, 1868) (Hyria) 
Eupithecia ferruginaria Moore, 1868. 
China, India, Sikkim. 

Hydrelia binotata Inoue, 1987 

China, Nepal. 

Hydrelia castaria (Leech, 1897) (Plemyria) 

Hydrelia cingulata Hampson, 1896 


Hydrelia condensata (Walker, 1862) 



Hydrelia conspicuaria (Leech, 1897) 



Remarks. This species and//, elegans (Inoue) were 
long assigned to Palpoctenidia. Both species were 
described from just a single female holotype. The 
discovery of a male specimen of conspicuaria has 
enabled us to examine the genitalia of that sex, which 
shows that it belongs to Hydrelia. Furthermore, there is 
a single small areole on the forewing in both 



conspicuaria and elegans, which is absent in most 
specimens of the only species of Palpoctenidia. 

Hydrelia controversa Inoue, 1982 


Hydrelia crocearia Hampson, 1896 


Hydrelia elegans (Inoue, 1982) comb. n. 


Hydrelia enisaria Prout, 1926 


Hydrelia flammeolaria (Hufnagel, 1767) 

Phalaena centrata Fabricius, [1776]. 
Asthena chibiana Matsumura, 1925. 
Geometra flavicata Thunberg, 1784. 
Phalaena flavostrigata Donovan, 1806. 
Geometra luteata [Denis & Schiffermuller], 1775 
Phalaena sinuosata Giorna, 1 79 1 

Widely distributed across the Palaearctic from western 
Europe to Japan. 

Hydrelia flammulata (Bastelberger, 1911) 



Hydrelia flavilinea (Warren, 1893) 


Sikkim, China. 

Hydrelia fuscocastanea Inoue, 1982 


Hydrelia gracilipennis Inoue, 1982 


Hydrelia impleta Prout, 1938 


Hydrelia inornata (Hulst, 1896) 

Tephroclystis inornata Hulst, 1896. 
Euchoeca exhumata Pearsall, 1906. 


Hydrelia laetivirga Prout, 1934 


Hydrelia latsaria (Oberthiir, 1893) (Acidalia) 

Hydrelia leucogramma Wehrli, 1931 


Hydrelia lineata (Warren, 1893) (Autallacta) 
Sikkim, China, Nepal. 

Hydrelia lucata (Guenee, [1858]) (Asthena) 

Hydrelia luteosparsata Sterneck, 1928 


Hydrelia marginepunctata Warren, 1893 

Sikkim, China, Nepal. 

Hydrelia microptera Inoue, 1987 

Nepal, China. 

Hydrelia musculata (Staudinger, 1897) 



Hydrelia nepalensis Inoue, 1987 

Nepal, China. 

Hydrelia nisaria (Christoph, 1881) (Acidalia) 
Hydrelia nisaria japonica Inoue, 1944 
Russia, China, Japan, Korea. 

Hydrelia ochrearia Leech, 1897 


Hydrelia ornata (Moore, 1868) (Hyria) 
India, Sikkim, Nepal, China. 

Hydrelia parvularia (Leech, 1897) (Plemyria) 

Hydrelia parvulata (Staudinger, 1897) 


Hydrelia pavonica Xue, 1999 


Hydrelia rhodoptera Hampson, 1895 

Sikkim, China. 

Hydrelia rubraria Hampson, 1903 

China (Tibet). 

Hydrelia rubricosta Inoue, 1892 

Nepal, China. 

Hydrelia rubrilinea Inoue, 1987 

Nepal, China. 

Hydrelia rubrivena Wileman, 1911 

China (Taiwan). 

Remarks. The abdomen of the type of rubraria 
Hampson is missing, so we have been unable to confirm 
the currently accepted identity as valid. H. rubrivena 
may be ajunior synonym of aurantiaca Hampson. The 
wing markings are similar but the colour differs 
between the two species. The male genitalia of both 
species are also quite similar. In the female, the corpus 
bursae is smaller in rubrivena and the signum shorter 
and broader. H. rubraria has the same markings as 
both rubrivena and aurantiaca. 

Hydrelia rufigrisea (Warren, 1893) 


Sikkim, China. 

Hydrelia rufinota Hampson, 1896 

India, China. 

Hydrelia sanguiflua Hampson, 1896 


Hydrelia sanguiniplaga Swinhoe, 1902 

China, Burma. 

Hydrelia scotozona Yazaki, 1995 


Hydrelia sericea sericea (Butler, 1880) 


China, Nepal, N. E. Himalaya. 

Hydrelia sericea pampesia Prout, 1938 


Hydrelia shioyana (Matsumura, 1927) 


Hydrelia adesma Prout, 1930. 

Hydrelia speciosa Inoue, 1992 


Hydrelia subcingulata Inoue, 1987 

Nepal, China. 

Hydrelia sublatsaria Wehrli, 1938 


Hydrelia subobliquaria (Moore, 1868) 


India, China, Nepal, Sikkim. 

Hydrelia subtestacea Inoue, 1982 

Nepal, China. 

Hydrelia sylvata ([Denis & Schiffermiiller], 

1775) (Geometra) 

Hydrelia sachalinensis Matsumura, 1925. 
Phalaena testaceata Donovan, 1810. 

Palaearctic, from western Europe to Japan. 

Hydrelia tenera (Staudinger, 1897) (Cidaria) 

Hydrelia terraenovae Krogerus, 1954 


Hydrelia ulula Bastelberger, 1911 

China (Taiwan). 

Hydrelia undularia (Leech, 1897) (Venusia) 
China, Nepal. 

Hydrelia undulosata (Moore, 1888) (Hyria) 

Species excluded 

Genitalia examined: flavidula (male, female). 

'Hydrelia' flavidula (Warren, 1907) (Hastina) 
Papua New Guinea. 

Remarks. This sexually dimorphic species does not 
belong to Hydrelia nor, indeed, to the Asthenini, 
although the uncus is absent. The frons is narrow and 
flat and the antenna in both sexes bears paired hair 
tufts, which are extremely long in the male. The labides 
are weak, united medially and bear a pair of setose 
processes. The valva is very long and narrow, with the 



costa strongly sclerotized and with a sharp terminal 
process; the sacculus lacks a process. The female 
genitalia are similar to those of Acolutha Warren in 
having a globose corpus bursae covered with 
Eupithecia-tike spines and an appendix bursae. 

Agnibesa Moore, 1888 

(Figs 69-75 ; 190; 198, 231; 271.) 

Agnibesa Moore, 1888: 256. Type species: Somatina 
pictaria Moore, 1868: 645. 

MOTH (Figs 69-75). Head. Frons almost as promi- 
nent as in Hydrelia, but less broad. Labial palpi minute. 
Male antenna ciliated. Wings. White, variously marked, 
all species with some yellow suffusion particularly on 
forewing. Venation as in Fig. 190. Forewing relatively 
elongated, with apex not pointed, termen smooth; areole 
single and small; vein R ( diverges from stem of R, s 
after (i.e., distal to) point of divergence of vein R . 
Hindwing with termen crenulated; discocellulars not 
biangulate; vein M, arising nearer M than M v 

Male genitalia (Figs 198,231). Labides expanded 
at apex. Juxta narrow to moderate. Valva: sacculus 
extended into short, narrow projection beyond the 
valva. Aedeagus: vesica bearing numerous minute 
denticles. Sternum A8 unmodified. 

Female genitalia (Fig. 27 1 ), Ductus bursae with 
strongly sclerotized antrum of varying length. Corpus 
bursaeglobose with scatteringofdenticlesorshort spines 
arranged as an ill-defined band across middle; signum 
composed of radiating denticles, sometimes with sec- 
ond, smaller signum nearer posterior end of corpus. 

Diagnosis. The adults of most species of Agnibesa 
are larger than in Hydrelia and always have a pure 
white ground colour. Whereas in Hydrelia vein R of 
the forewing diverges from the common stem before 
R 5 diverges, in Agnibesa R ( diverges after (distal to) 
the divergence point of R,. The vesica of the aedeagus 
bears many minute denticles in Agnibesa but not in 
Hydrelia. The corpus bursae in Agnibesa has a 
conspicuous, largely medial scattering of denticles, 
which are lacking in Hydrelia. Agnibesa is 
distinguished from Euchoeca by the fact that the moths 
are larger, by the narrower forewing, the white ground 
colour and the presence of minute denticles on the 

Distribution. China, India, Nepal, Sikkim. 

Species included. Six species. Genitalia examined: 
pictaria pictaria (male, female), pictaria brevibasis 
(male), pleopictaria (male, female), plumbeolineata 
(male, female), punctilinearia (male, female), 
recurvilineata recurvilineata (male, female), 
recurvilineata mewplyta (male, female), venusta 

Agnibesa pictaria pictaria (Moore, 1868) 

(Somatina ?) 

India, Nepal, Sikkim, China (Tibet, from the frontier to 
Nepal to Medog). 

Agnibesa pictaria brevibasis Prout, 1938 

China (Shanxi, Gansu, Sichuan, Yunnan, Tibet 
(recorded at Bomi, which is N E of Medog)). 

Remarks. The two subspecies of pictaria are 
geographically separated in Tibet by Mt Namjagbarwa 
(Xue&Zhu, 1999). 

Agnibesa pleopictaria Xue, 1999 


Agnibesa plumbeolineata (Hampson, 1895) 


Sikkim, China. 

Agnibesa punctilinearia (Leech, 1897) 



Agnibesa recurvilineata recurvilineata 
Moore, 1888 

India, Nepal, Sikkim. 

Agnibesa recurvilineata meroplyta Prout, 


Agnibesa venusta Warren, 1897 

Sikkim, Nepal. China. 

Euchoeca Hiibner, [1823] 1816 

(Figs 76; 199,232:272.) 

Euchoeca Hiibner, [1823] 1816: 298. Type species: 
Geometra hepararia Hiibner, [1799] 1796: pi. 11, 
fig. 58, a junior synonym of £. nebulata (Scopoli, 

Moth (Fig. 76). Head. Frons a little less protuberant 
than in Hydrelia. Labial palpi minute, hardly extend- 
ing beyond frons. Antenna with short cilia. Wings. 
Grey-brown. Forewing almost triangular, costa almost 
straight, termen weakly convex; areole single, vein R ; 
stalked with R.,, diverging from stem after R 5 , which 
diverges at apex of areole (as in Agnibesa). Hindwing: 
termen smooth, angled at position of vein M ; 
discocellulars not biangulate; vein M 2 arising well 
anterior to middle of discocellulars. 



Male genitalia (Figs 199, 232). Saccus broad. 
Labides narrow, curved, extending beyond tegumen. 
Juxta in form of a broad plate extending into a slender 
digitate process. Valva narrow at base, broadening 
apically into a broader lobe; sacculus extending just 
beyond margin of valva. Aedeagus: vesica lacking 
cornuti. Sternum A8 unmodified. 

Female genitalia (Fig. 272). Ductus bursae weakly 
sclerotized. Corpus bursae with two signa and a 
scattering of minute denticles; signum prominent, long 
and narrow, composed of radiating denticles; small, 
approximately round denticulate signum situated 

DIAGNOSIS. Unlike the condition in the forewing of 
Hydrelia, in Euchoeca nebulata vein R { diverges distal 
to the divergence point of vein R-. The apex of the 
forewing is more rounded than in most species of 
Hydrelia. The male genitalia exhibit some similarity to 
those of Agnibesa and Asthenotricha, but E. nebulata 
is distinctly smaller than in these other genera and 
differs also in being almost uniformly grey-brown. 
The scattered denticles on the corpus bursae of 
Euchoeca are minute and far less prominent than those 
found in Agnibesa or Asthenotricha. 

Remarks. Only one species is recorded under this 
genus. The species, cichisa Prout, 1939 (Prout, 1934- 
1939: 253, pi. 18: c, type specimens from West China: 
Mt Omei, Gipfel., in BMNH, examined) was placed 
under Euchoeca by Parsons et al. (in Scoble, 1999) 
under the mistaken assumption that Prout had described 
it under that genus. In fact, cichisa was originally 
described under Eupithecia where it properly belongs. 

Distribution. Europe, Transcaucasia, Russia, 

SPECIES INCLUDED. The genus is monotypic. Geni- 
talia examined: nebulata (male, female). 

Euchoeca nebulata (Scopoli, 1763) 


Geometra hepararia Hiibner, [1799]. 

Geometra heparata [Denis & Schiffermuller], 1775 

Phalaena obliterata Hufnagel, 1767 

Phalaena strigata Thunberg, 1 788 

Widespread in Europe, and across Russia to Japan. 

FOODPLANTS. Betulaceae: Alnus glutinosa; Alnus 
incana; Alnus. 

Epicyme Meyrick, 1885 

(Figs 77; 200, 233; 273.) 

Epicyme Meyrick, 1885: 589. Type species: 
Ptychopoda rubropunctaria Doubleday, 1843: 287. 
[Replacement name for Hippolyte Meyrick.] 

Hippolyte Meyrick, 1883: 526. Type species: 
Ptychopoda rubropunctaria Doubleday, 1843. [Jun- 
ior homonym of Hippolyte Leach, [1814] 1830 

MOTH (Fig. 77). Head. Frons narrow, not prominent. 
Labial palpi narrow and short. Antenna simple, ciliated 
in male. Wings. Brownish with numerous wavy trans- 
verse lines, female with large, darker brown, irregular 
spots on lower half of forewing at postmedial position. 
Wing shape and venation of forewing as in Hydrelia, 
but vein R p R M and R 5 arising independently from 
near apex of areole, not sharing a common stem. 
Hindwing: apex pointed, termen weakly angled medi- 
ally; vein M 3 of the hindwing stalked with CuA r 

Male genitalia (Figs 200, 233). Labides present 
as membranous outpushings of diaphragmata (diffi- 
cult to discern in slide preparations) with numerous 
setae. Juxta with narrow extension. Valva weakly but 
distinctively curved; hair-like scales arising from apex; 
costa weakly sclerotized, broadened slightly at middle; 
sacculus not extended or otherwise modified. 
Aedeagus: vesica lacking cornuti. 

FEMALEGENlTALlA(Fig.273). Ductus bursae mostly 
membranous, anterior portion sclerotized. Corpus 
bursae with small bundle of spines near cervix bursae; 
signum elliptical, composed of long radiating spines. 

Diagnosis. Many external characters are similar to 
those in Hydrelia, but the frons is narrow and not 
prominent in Epicyme rubropunctaria. The genitalia, 
however, differ strongly: in Epicyme the valva bears a 
series of long, fixed hair-like setae with expanded 
apices around its distal margin, the sacculus of the 
valva is not sclerotized and lacks an extension, the 
ductus bursae is unsclerotized posteriorly, and the 
signum is composed of long radiating spines rather 
than denticles. The male genitalia of E. rubropunctaria 
are similar to those of the monotypic Peruvian genus 
Leucoctenorrhoe, but there are many differences 
between them: notably, in Leucotenorrhoe the antenna 
are bipectinate (in both sexes); vein R, of the forewing 
arises independently from near the apex of the areole; 
and vein M, of the hindwing is stalked with CuA . 

Distribution. Australia, New Zealand. 

Remarks. Unlike most Asthenini, the frons of this 
species is not prominent, the sacculus of the valva is 
not extended, and the labides are inconspicuous, mem- 
branous outpushings. However, the reduced condition 
of the labial palpi and the uncus, and the signum with 
its radiating spines means that the genus is reasonably 
included in the tribe. 

Species INCLUDED. One species is known. Genitalia 
examined: rubropunctaria (male, female). 


Epicyme rubropunctaria (Doubleday, 1843) 

Asthena mullata Guenee, 1 868. 
Aslhena risata Guenee, [ 1 858]. 
Asthena vexata Walker, 1869. 

Australia, New Zealand. 

Eschatarchia Warren, 1894 

(Figs 78; 201, 234; 274.) 

Eschatarchia Warren, 1894: 395. Type species: 
Eschatarchia lineata Warren, 1894: 295. 

MOTH (Fig. 78). Head. Frons not broadened, slightly 
protuberant. Labial palpi minute, projecting slightly 
beyond head. Antenna in both sexes ciliate. Wings 
broad, off-white with strong markings at termen; both 
wings with termen angled at M, and crenulated from 
apex to angle, nearly straight below angle; vein R ( 
diverging from the common stem of R,^ beyond (distal 
to) the divergence point of R s ; venation almost identi- 
cal to that of Agnibesa, but forewing with areole larger. 

Male genitalia (Figs 201, 234). Saccus large. 
Labides broad, as long as the length of tegumen, 
expanded terminally into relatively membranous head. 
Juxta slightly constricted medially. Valva broad; 
sacculus well sclerotized, extending beyond margin, 
asymmetrical between each valva in shape and size. 
Aedeagus: vesica with patch of cornuti and three 
sclerotized plates. Sternum A8 of abdomen unmodi- 

Female GENITALIA (Fig. 274). Apophyses anteriores 
short. Ductus bursae strongly sclerotized to the distal 
part of corpus bursae. Corpus bursae covered almost 
entirely with minute denticles; signum composed of 
radiating denticles. 

Diagnosis. The wing colour and pattern of 
Eschatarchia lineata resembles that of Chalyboclydon 
marginata (excluded from Asthenini, see p. 106). The 
asymmetrical valvae and the wing markings are good 
distinguishing features of Eschatarchia. It is further 
distinguished from Hydrelia by its forewing venation, 
vein R : diverging from the common stem of R 
beyond (distal to) the divergence point of R_. 

Distribution. Japan, China, Burma. 

Species included. One described species. Genita- 
lia examined: lineata fornwsana (male), lineata lineata 
(male, female), undescribed species from Burma and 
West China (male). 

Eschatarchia lineata Warren, 1894 

Hydrelia angularia Leech, 1897 

Eschatarchia lineata fornwsana Inoue, 1970. Syn. n. 


Japan, China, Burma. 

Remarks. The subspecies /bwjoKwa was described 
(Inoue, 1970) because of its more deeply incurved 
submarginal dark line on the forewing and the slightly 
more pronounced angle of the termen. These characters, 
however, can be found in certain specimens of the 
nominate subspecies from Japan, and some of them 
vary even more strongly. The male genitalia of the type 
specimens of lineata and fornwsana are indistinguish- 

Prout (1934-1939: 180) wrote of a darker 'form' of 
lineata from Burma. The median band on the under- 
side of both wings, which is composed of four dark 
lines, is complete, not interrupted or reduced. The male 
genitalia also differ somewhat from lineata: the saccu- 
lus lobe of the left side is markedly narrower, and the 
main bundle of cornuti are very weak and small, with 
the individual spines much shorter. Prout predicted 
that the form would probably be discovered in West 
China. We have, indeed, located a male from Sichuan 
Province, West China, which is quite similar to the 
specimen from Burma. Given the differences, it seems 
likely that the form from Burma and West China 
represents an undescribed species. 

'Chalyboclydon 'flexilinea Warren, 1898 

(Figs 79; 202, 235:275.) 

The generic placement of flexilinea is unclear. It is not 
congeneric with the type species of Chalyboclydon 
Warren (C. marginata Warren), with which it was 
combined in the past. However, given the substantial 
amount of work that is evidently needed on the 
Larentiinae, we prefer at this stage neither to force a 
doubtful recombination with another asthenine genus, 
nor to describe a new genus for flexilinea. Details of 
marginata are provided under Chalyboclydon below. 
The wing markings of flexilinea and marginata are 
similar but distinct (compare Figs 79, 1 80); the venation 
and genitalia differ markedly. 

Moth (Fig. 79). Head. Frons slightly more promi- 
nent than in marginata, labial palpi shorter. Antenna 
simple in both sexes. Wings. Forewing: apex less 
produced, termen hardly marked. Hindwing with hind 
margin elongate and a little incurved, anal angle 
pointed; termen hardly angled. Venation: forewing 
with areole absent; stalk of R, 5 arising from upper 
angle of cell; R 1 diverges from common stalk after 
(distal to) point of divergence of R ? ; hindwing with 
Sc+R ( combined with Rs to near end of cell, Rs stalked 
with M ; cell in both wings extremely short, maximum 
length less than 1/3 of wing length; discocellulars less 
oblique than in marginata; M arising slightly closer to 
M, than to M 3 ; M, stalked with CuA |; CuA 2 arising 
near lower angle of cell. 



Male genitalia (Figs 202, 235). Saccus broad, 
with a narrower, nearly quadrate extension. Labides 
well developed, with bases triangular, apex slightly 
broadened and setose. Juxta broad. Valva costa simple; 
sacculus much simpler than in marginata, with small 
spine-like process extending just beyond margin of 
valva. Aedeagus: vesica with some scattered denticles. 
Abdomen: sternum A8 unmodified. 

Female genitalia (Fig. 275). Ductus bursae long, 
anterior half sclerotized. Corpus bursae long, shorter 
than ductus bursae, with two small signa composed of 
radiating denticles, one near cervix bursae and another 
at middle, dense covering of denticles on side opposite 
signa. Sternum A7 not modified. 

DIAGNOSIS. ' C. 'flexilinea differs from C marginata 
in several ways. The forewing lacks an areole and in 
the hind wing M is stalked with CuA,. The very short 
cell in both wings differs from almost all other genera 
other than Cleptocosmia. In Cleptocosmia the palpi 
are longer and the basal half of the forewing is clothed 
with erect furry hair in the male. The male and female 
genitalia are distinctive. 

Distribution. India, Sikkim, Burma. 

Genitalia examined: flexilinea (male, female). 

'Chalyboclydon'flexilinea Warren, 1898: 22 

India, Sikkim, Burma. 

Remarks. The species was described from a single 
female specimen, labelled as flexilinea by Warren and, 
incorrectly, as Chalyboclydon marginata Warren by 
Prout (for details see below under Chalyboclydon). 

Palpoctenidia Prout, 1930 

(Figs 80, 81; 203, 236; 259; 276.) 

Palpoctenidia Prout, 1930: 311. Type species: 
Chrysocraspeda phoenicosoma Swinhoe, 1895: 294. 

Moth (Figs 80,8 1 ). Head. Frons broad, moderately 
protuberant in male, flat in female. Labial palpi slender, 
extending slightly beyond frons. Antenna bipectinate 
in male, simple in female. Wings broad; forewing grey- 
brown with broad postmedian band nearly reaching 
termen; apex slightly produced, termen gently rounded, 
not angled at middle; areole absent in most specimens; 
vein R : diverging from stem of R before (proximal 
to) point of divergence of R 5 ; M arising from cell, 
which is nearly as long as half length of forewing. 
Hindwing with termen angled at middle; vein Sc+R t 
combined with Rs to 3/4 of cell; veins Rs and M 
sharing short stalk; discocellulars not biangulate; vein 
M 2 arising well anterior to middle of discocellulars. 

Male genitalia (Figs 203, 236, 259). Saccus broad, 
weakly excavated. Labides short, curved, well- 

sclerotized with apices swollen and united medially, 
each element with small teeth projecting ventrally. 
Juxta: narrowing towards apex. Valva: densely setose 
at apex; sacculus distinctive, excavated terminally so 
appearing double pointed, with several strong spines 
subapically; base with long hairs arising from mem- 
branous pouch on each side. Aedeagus weakly curved; 
vesica lacking cornuti. Sternum A8 (Fig. 259) bearing 
row of setae along distal margin, otherwise unmodi- 

Female genitalia (Fig. 276). Bursa copulatrix: 
ductus bursae short, sclerotized throughout; corpus 
bursae globose, signum long, narrow and composed of 
stout radiating spines, arranged more sparsely than in 
usual condition, and diminishing in size to apex; surface 
of corpus with scattered minute denticles. Segment A7 
with pair of lateral pockets. 

Diagnosis. The strong subapical spines on the 
sacculus and the row of setae along the distal margin of 
the abdominal sternum A8 in the male distinguish the 
single species of Palpoctenidia from all other genera 
in the Asthenini. Palpoctenidia resembles 
' Chalyboclydon flexilinea in lacking an areole on the 
forewing and by the similar wing markings, notably 
the strongly protuberant postmedian band on the 
forewing. This band almost touches the termen medi- 
ally whereas in flexilinea the band actually meets the 
termen. Palpoctenidia is also clearly distinguished 
from flexilinea by the bipectinate condition of the male 
antenna and the marked differences in the genitalia. 

Remarks. The excavated sacculus is similar to that 
found in Asthena. Although an areole is absent in most 
material of P. phoenicosoma, in four specimens in the 
BMNH from 'Rantaizan, Formosa' [Taiwan], collected 
by Wileman a single small areole is present. 

Asthenine features of Palpoctenidia include the ab- 
sence of an uncus and the presence of typically 
asthenine labides. The extension of the sacculus is 
notable and much modified. Although the signum is 
composed of stout spines, their radiating arrangement 
is merely a modification of the basic asthenine plan. 

Distribution. China, Japan, India. 

SPECIES INCLUDED. One species. Genitalia examined: 
phoenicosoma phoenicosoma (male, female), 
p. semilauta (male). 

Palpoctenidia phoenicosoma phoenicosoma 
(Swinhoe, 1895) (Chrysocraspeda) 

India, China. 

Palpoctenidia phoenicosoma semilauta Prout, 



Asthenotricha Warren, 1899 

(Figs 82-1 1 1; 204, 205; 237, 238; 277, 278.) 

Asthenotricha Warren, 1899: 34. Type species: 
Asthenotricha dentatissima Warren, 1899: 34. 

Astenotricha Debauche, 1938: 40. (An incorrect 
subsequent spelling.) 

Moth (Figs 82-1 1 1). Head. Frons less prominent 
than in Hydrelia. Labial palpi stronger and longer than 
in typical Hydrelia. Antenna: ciliated in male. Wings. 
Ochreous to purplish brown, often with broad fascia 
across forewings and hindwings forming a U in resting 
posture. Forewing: fairly broad; termen rounded, apex 
often pointed; tornus rounded; areole single; vein R : 
diverges from R,^ proximal to point at which R s 
diverges; male of dentatissima group with patch of 
modified scales on dorsal and ventral surfaces. 
Hindwing: relatively broad, termen rounded; male of 
dentatissima group with costa markedly expanded and 
with large hairpencil on dorsal surface; hindwing un- 
modified in argyridia group; discocellulars not 
biangulate; anastomosis of Sc+Rj with Rs rather shorter 
than in most Larentiinae; vein Rs not stalked with M 
in type species, but stalked in most species. 

MALE GENITALIA (Figs 204, 205, 237, 238). Tegumen/ 
?uncus with distinctive, long, narrow anterior exten- 
sion. Saccus broad, short. Labides in form of a pair of 
finger-like processes, usually extending to just beyond 
middle of tegumen. Juxta W-shaped, often with central 
arm long. Valva: costa straight; sacculus with well- 
developed finger-like extension. Aedeagus: vesica 
simple, without cornuti. 

Female genitalia (Fig. 277, 278). Ductus bursae: 
antrum with very wide mouth, narrowing so that antrum 
appears conspicuously Y- or V-shaped; colliculum 
usually very short and situated distinctively in middle 
of the membranous duct. Corpus bursae: signum long, 
narrow or pear-shaped, composed of radiating spines; 
prominent (Eupithecia-Wke) denticles present or absent 
from corpus bursae. 

DIAGNOSIS. In Asthenotricha the most notable 
distinguishing features are the extended uncus/ 
tegumen, the W-shaped juxta, the Y- or V-shaped antrum 
and the very short colliculum situated, usually, in the 
middle of the ductus bursae. The hindwing is broader 
than in Hydrelia. even in those species where it is not 
markedly expanded in the male. 

REMARKS. We have included Asthenotricha in the 
Asthenini despite the presence of well developed labial 
palpi. What appears to be an uncus is also stronger than 
is typical for the tribe, and there are Eupithecia-Mke 
denticles on the corpus bursae of some species. How- 
ever, the sacculus is extended, the form of the labides 
fits the condition widely encountered in the tribe, and 


the signum is typically asthenine, being composed of 
radiating spines. 

Distribution. Afrotropics, including Madagascar. 

Species included. Thirty-four. 

The dentatissima group 

This group is distinguished by the presence of a hair 
tuft on the hindwing of the male and a broadened costa. 

Species included. Twenty-six. Genitalia examined: 
ambly coma (male), anisobapta (male, female), barnsae 
(male, female), dentatissima (male, female), flavicoma 
(male), lophopterata ( male, female), malostigma (male, 
female), polydora (male, female), proschora (male, 
female), pycnoconia (male), serraticornis (male, 
female), torata (male, female), tripogonias (male, 

Asthenotricha amblycoma Prout, 1935 

Equatorial Guinea. 

Asthenotricha anisobapta Prout, 1932 

Kenya, Uganda. 

Asthenotricha ansorgei Warren, 1899 

Kenya. Uganda. 

Asthenotricha barnsae Prout, 1935 

Kenya, Uganda. 

Asthenotricha comosissima Herbulot, 1970 


Asthenotricha deficiens Herbulot, 1954 


Asthenotricha dentatissima Warren, 1899 

Tanzania. Kenya, Uganda, Zaire. 

Asthenotricha fernandi Prout, 1935 

Equatorial Guinea. 

Asthenotricha flavicoma Warren, 1899 

Cameroon, Uganda, Zaire. 

Asthenotricha furtiva Herbulot, 1960 


Asthenotricha grandis Herbulot, 1997 




Asthenotricha lophopterata (Guenee, [1858]) 


Madagascar, Reunion. 

Asthenotricha malostigma Prout, 1921 


Asthenotricha nesiotes Herbulot, 1954 


Asthenotricha paraholica Herbulot, 1954 


Asthenotricha polydora Debauche, 1938 (as 


Uganda, Zaire. 

Asthenotricha proschora Fletcher, 1958 

Uganda, Zaire. 

Asthenotricha psephotaenia Prout, 1935 

Uganda, Zaire. 

Asthenotricha pycnoconia Janse, 1933 

South Africa, Uganda. 

Asthenotricha pythia Debauche, 1938 (as 



Asthenotricha quadrata Herbulot, 1960 


Remarks. In this species the hind margin of the 
forewing is not elongate. Both wings are shaped in a 
similar way to those in Hydrelia. 

Asthenotricha semidivisa semidivisa Warren, 

Cameroon, Uganda. 

Asthenotricha semidivisa euchroma Prout, 


Asthenotricha serraticornis Warren, 1902 

Kenya, Tanzania, Uganda. 

Asthenotricha straba Prout, 1921 

Angola, Cameroon, Kenya, Uganda, Zaire. 

Asthenotricha torata Prout, 1932 


Remarks. This species differs from other species of 
the genus, as does tripogonias, by the forewing of the 
male, which is folded and bears conspicuous 
androconia. Prout (1932: 108) expressed his doubts 
that the species belonged to Asthenotricha. However, 
in both species the male genitalia fit the general pattern 
exhibited by the type species. Although the corpus 
bursae bears Eupithecia-Mke denticles, the antrum and 
colliculum are characteristic of Asthenotricha. Females 
differ from males in colour and markings. Contrary to 
Prout's description, a hair-tuft is in fact present on the 
hindwing of males of torata, although it is smaller and 
paler than in other species. 

Asthenotricha tripogonias Prout, 1926 


The argyridia group 

In argyridia and its relatives, the hair tuft on the 
hindwing of the male is lacking and the costa of the 
hindwing is not broadened. 

Species included. Eight species. Genitalia exam- 
ined: argyridia (male, female), costalis (male), inutilis 
(male), meruana (male), sjostedti sjostedti (male), 
strangulata (male), unipecten (male, female). 

Asthenotricha argyridia (Butler, 1894) 
comb. n. (Cataclysme) 

Eulype? disparata Warren, 1897 

Kenya, Rwanda, Uganda. 

Asthenotricha candace (Prout, 1929) 
comb. n. (Hydrelia) 


Asthenotricha costalis (Aurivillius, 1910) 
comb. n. (Hydrelia) 


Asthenotricha inutilis Warren, 1901 

Cameroon, Kenya, South Africa, Uganda. 

Asthenotricha meruana (Aurivillius, 1910) 
comb. n. (Hydrelia) 


Asthenotricha sjostedti sjostedti (Aurivillius, 
1910) comb. n. 

Hydrelia sjostedti Aurivillius, 1910. 

Kenya, Tanzania. 



Asthenotricha sjostedti mionoseista (Prout, 
1921) comb. n. (Hydrelia) 

Rwanda, Zaire. 

Asthenotricha strangulata Herbulot, 1953 


Asthenotricha unipecten (Prout, 1915) 


Hydrelia unipecten tamsi Prout, 1935. 

Kenya, Uganda, Zaire. Sao Tome & Principe. 

Remarks. Prout (1915; 1934-1939) noted that the 
antenna of the male is unipectinate. In fact this condition 
also occurs in the female, although the pectinations are 
shorter in this sex. Unipectinate antennae are rare in 
Larentiinae so this is an unusual character. 

Further study is required to assess the taxonomic 
status of Prout's subspecies tamsi. 

Venusia Curtis, 1839 

(Figs 1 12-140; 191; 206-208; 239-241; 279-280.) 

Venusia Curtis, 1839: 759. Type species: Venusia 

cambrica Curtis, 1839: 759. 
Discoloxia Warren, 1895: 105. Type species: Cidaria 

obliquisigna Moore, 1888: 278. 

Moth (Figs 112-140). Head. Frons broad, gener- 
ally protuberant. Labial palpi minute. Antenna of male 
ciliate or bipectinate with pectinations short. Wings. 
Whitish to grey; forewing usually darker than hindwing; 
rarely uniformly brown; broad, termen of both wings 
gently curved, not angled or dentate. Venation as in 
Fig. 191). Forewing with a single areole, R : and R, 
arising from apex of areole, or R : diverging after short 
union with R, ; vein M : diverging from areole; 
discocellulars curved and oblique, occasionally 
biangulate. Hindwing with discocellulars biangulate, 
vein M, arising from the second angle, close to M . 

Male genitalia (Fig. 206-208; 239-241). Saccus 
generally broader and longer than tegumen. Labides 
extending from bases of valvae with slightly expanded, 
setose apices, bases united with those of transtillae. 
Juxta weakly sclerotized, plate-like, sometimes pointed 
posteriorly. Valva: usually narrowing to apex rather 
than rounded; sacculus with prominent extension other 
than in dilecta Yazaki where process is absent. 
Aedeagus simple, vesica lacking cornuti. Sternum A8 
of male not modified. 

Female genitalia (Figs 279, 280). Ductus bursae 
weakly sclerotized throughout length other than for a 
short membrane. Corpus bursae generally globose, 
often with numerous minute denticles, sometimes 
dense, or small spines; signum usually long and narrow, 

sometimes pear-shaped, sometimes small and almost 
round, composed of radiating denticles, occasionally 
with additional small signum distally. 

Diagnosis. Similar to Hydrelia, but the hindwing 
discocellulars of Venusia are distinctly biangulate and 
sometimes the male antennae are bipectinate. In the 
female of Venusia, the surface of the corpus bursae 
lacks denticles more often than in Hydrelia. In those 
species of Venusia where denticles are present, they 
tend to be arranged more densely. 

Distribution. Palaearctic, Nearctic and Oriental 

Species included. Forty-two described species, one 
doubtfully included. Genitalia examined: albinea 
(male, female), apicistrigaria(femalt), balausta (male, 
female), biangulata (male, female), blame ri blame ri 
(male, female), cambrica (male, female), comptaria 
(male, female), conisaria conisaria (male, female), 
crassisigna (male, female), eucosma (male), 
kioudjrouaria (female), laria laria (male, female), 
lilacina lilacina (male, female), limata (male, female). 
maniata (male, female), marmoraria (male, female), 
naparia (male, female), nigrifurca (male, female), 
obliquisigna (male, female), ochrota (mii\e),pallidaria 
(male, female), paradoxa (male), pearsalli (male, 
female), planicaput (male, female), punctiuncula 
(male, female), purpuraria (male, female), roseicosta 
(male), scitularia (male, female), scitularia (male, 
female), sikkimensis (male, female), tchraria (male, 
female), violettaria (male, female), yasudai (male, 

Venusia accentuata (Prout, 1914) 


Venusia albinea (Prout, 1938) (Discoloxia) 


REMARKS. The female genitalia of this species has a 
small, almost round signum composed of radiating 
denticles. Otherwise albinea is a typical species of 

Venusia apicistrigaria (Djakonov, 1936) 



Venusia balausta Xue, 1999 


Venusia biangulata (Sterneck, 1938) 




Venusia blomeri blomeri (Curtis, 1832) 

Cidaria pulchraria Eversmann, 1842. 

Widespread in Europe, across Estonia, Latvia, Lithua- 
nia, Russia to Japan, China (far N E), and North 

Foodplants. Ulmaceae: Ulmus glabra; Ulmus. 

Venusia blomeri euchloe (Bryk, 1949) 



Venusia blomeri szechuanensis Wehrli, 1931 

China (S W). 

Venusia brevipectinata Prout, 1938 


Venusia cambrica cambrica Curtis, 1839 

Venusia cambrica shaotsii Bryk, 1949. 

Venusia cambricaria Guenee, [1858] [Emendation of 
cambrica Curtis.] 

Hydrelia cambricata Herrich-Schaffer, 1861. [Emen- 
dation of cambrica Curtis.] 

Eubolia erutaria Boisduval, 1 840 

Acidalia nebulosaria Freyer, 1 850 

Tephrosia scitularia Walker, 1 860 

Widespread across the Holarctic region. 

Foodplants. Betulaceae: Alnus incana tenuifolia; 
Alnus rubra: Betula; Ericaceae: Vaccinium; Rosaceae: 
Amelanchier; Malus; Sorbus americana; Sorbus 
aucuparia; Sorbus; Salicaceae: Populus trichocarpa; 

Venusia cambrica aphrodite Bryk, 1942 

Russia (Kurile Islands). 

Venusia comptaria (Walker, 1860) (Tephrosia) 

Venusia palumbes Franclemont, 1938. 
Larentia perlineata Packard, 1873. 
Euchoeca salienta Pearsall, 1905. 

Canada, U.S.A. 

Venusia conisaria conisaria Hampson, 1903 

China, Nepal, Sikkim. 

Venusia conisaria hypoconia (Prout, 1938) 



Venusia crassisigna Inoue, 1987 

Nepal, China. 

Venusia dilecta Yazaki, 1995 


Venusia eucosma (Prout, 1914) (Discoloxia) 

Venusia inefficax (Prout, 1938) (Discoloxia) 

Venusia kasyata Wiltshire, 1966 


Venusia kioudjrouaria Oberthiir, 1893 


Venusia laria laria Oberthiir, 1893 

Venusia laria Oberthiir, 1893. 

Venusia laria ilara (Prout, 1938) (Discoloxia) 

Venusia lilacina lilacina (Warren, 1893) 


Sikkim, Nepal. 

Venusia lilacina melanogramma Wehrli, 1931 
subsp. rev. (Venusia (Discoloxia)) 


Venusia lilacina rala (Prout, 1938) 



Venusia limata Inoue, 1982 


Venusia lineata Wileman, 1916 


Venusia maniata Xue, 1999 


Venusia marmoraria (Leech, 1897) (Hydrelia) 



Venusia megaspilata (Warren, 1895) 



Venusia naparia Oberthur, 1893 


Venusia nigrifurca (Prout, 1926) (Discoloxia) 
Burma, China. 

Venusia obliquisigna (Moore, 1888) (Cidaria) 

India, Nepal, China. 

Venusia ochrota Hampson, 1903 

Venusia roseicosta Yazaki, 1994, syn. n. 
China, Nepal. 

Venusia pallidaria Hampson, 1903 


Venusia paradoxa Xue, 1999 


Venusia participata (Saute r, 1869) 



Venusia pearsalli (Dyar, 1906) (Euchoeca) 

Canada, U.S.A. 

FOODPLANTS. Aceraceae: Acer circinatum; 
Betulaceae.: Alnus incana tenuifolia.Alnus rubra; Abuts 
viridis sinuata; Betula; Cornaceae: Comus nuttalli; 
Fagaceae: Quercus garryana; Rosaceae: Crataegus; 
Malus; Salicaceae: Populus tremuloides; Populus 
trichocarpa; Salix. 

Venusia phasma (Butler, 1879) (Emmelesia) 

Venusia planicaput Inoue, 1987 

Nepal, China. 

Venusia punctiuncula Prout, 1938 


Venusia purpuraria (Hampson, 1895) 


Venusia scitula Xue, 1999 


Venusia semistrigata semistrigata (Christoph, 
1881) (Cidaria) 


Venusia semistrigata expressa Inoue, 1963 


Venusia sikkimensis (Warren, 1893) 


Sikkim, Bhutan, Nepal, India, China. 

Venusia syngenes Wehrli, 1931 (Venusia 


Venusia tchraria Oberthur, 1893 


Venusia violettaria Wehrli, 1931 (Venusia 

Venusia (Discoloxia) violettaria kukunoora Wehrli, 


Venusia yasudai Inoue, 1987 


Species of uncertain identity 

'Venusia' distrigaria (Boisduval, 1833) 


Remarks. Boisduval (1833: 263) wrote of this 
species, under the name Geometra distrigaria 
Boisduval 'Elle a le port de la Bilinearia [sicj d'Europe, 
mais elle est un tiers plus petite.' However, there were 
no Geometridae species from Europe (or from any- 
where else) named 'bilinearia' before 1833. In his 
checklist to the Lepidoptera of Madagascar, Viette 
(1990: 248) cited distrigaria as unidentified and with- 
out the type having been found. Although the species 
was placed in Venusia by Parsons et al. (in Scoble, 
1999), and in the card index to Geometridae in the 
BMNH, its identity and generic placement remain 

Nomenia Pearsall, 1905 gen. rev. 

(Figs 141;209, 242:281.) 

Nomenia Pearsall, 1905: 126. Type species: Larentia 

duodecimlineata Packard, 1873: 19. 
Nomenia Pearsall; Ferguson in Hodges et al., 1983: 

104. (Listed as a junior synonym of Venusia Curtis.) 



Moth (Fig. 141). Head. Frons weakly protuberant. 
Labial palpi short and slender. Antenna: male with a 
short, single, fasciculate protrusion on each 
flagellomere; female simple. Wings. Usually with bands 
of multiple transverse lines, sometimes lines not 
conspicuous; termen smooth. Forewing medium grey; 
vein R ( stalked with R M , R 5 arising independently 
from apex of areole, or, rarely, stalked with stem of 
R ; areole single; discocellulars strongly biangulate. 
Hindwing: pale grey to almost white, distinctly paler 
than forewing; discocellulars biangulate. 

Male genitalia (Figs 209, 242). Saccus smaller 
than tegumen. Labides extending to apex of tegumen, 
densely setose laterally. Juxta flask-shaped. Valva: 
costa densely setose, margin weakly convex; sacculus 
with long, finger-like extension, and a small, spined 
hump at base of extension. Aedeagus simple, without 
cornuti. Sternum A8 unmodified. 

Female genitalia (Fig. 28 1 ). Bursa copulatrix: duc- 
tus bursae sclerotized throughout; corpus bursae lacking 
signum, covered with Eupithecia-like spines except 
posteriorly, spines smaller towards anterior end of 
corpus bursae. 

DIAGNOSIS. The genitalia of Nomenia exhibit a 
general similarity to Venusia, but the genus differs in 
the presence of a single protrusion on each flagellomere 
in the male and the presence of setae on the labides. 
The presence of numerous Eupithecia-type spines on 
the corpus bursae of the female of Nomenia and the 
absence in that sex of a discrete signum clearly distin- 
guishes it from Venusia. 

REMARKS. The similarities between the male genita- 
lia of Nomenia and Venusia are notable. However, we 
prefer to revive Nomenia as a separate genus particu- 
larly because of the absence of a signum and the 
presence of typical Eupithecia-style denticles on the 
corpus bursae. Despite the absence of a typical 
asthenine signum, we retain the genus within the tribe 
given the close similarity between the genitalia and 
those of Venusia, and because of the reduced labial 
palpi. The absence of the asthenine signum is, there- 
fore, interpreted as a loss. 

Distribution. North America. 

Species included. Two species. Genitalia exam- 
ined: duodecimlineata duodeeimlineata (male, female). 

Nomenia duodecimlineata duodecimlineata 
(Packard, 1873) (Larentia) 

Nomenia unipecta Pearsall, 1906 
U.S.A. Type locality: California. 

Nomenia duodecimlineata secunda Pearsall, 

U.S.A.: Type locality: Colorado. 

Remarks. Listed as a subspecies by Ferguson in 
Hodges etai, 1983: 104. 

Nomenia obsoleta Swett, 1916 


Hastina Moore, 1888 

(Figs 142, 143; 192; 210, 243; 282.) 

Hastina Moore, 1888: 260. Type species: Hastina 
caeruleolineata Moore, 1888: 260. 

Moth (Fig. 142, 143). Head. Frons rounded, 
prominent. Labial palpi reduced, not significantly 
extended beyond frons. Antenna simple in both sexes, 
ciliated. Wings. Dark brown with weak yellowish lines 
(caeruleolineata) or pale brown with conspicuous and 
wider yellow lines (pluristrigata). Venation as in Fig. 
192. Forewing: broad, apex pointed, termen concave 
below apex, produced at the end of M and CuA r 
straight below CuA, ; areole single, small, vein R t long, 
stalked with R vein R 5 not stalked or only very 
briefly stalked with R H , R l diverging from the stem 
well distal to base of R . Hindwing: strongly crenulated, 
with sharp 'teeth' at the end of M 1 and M and excised 
between them; cell short, discocellulars angled at 
middle, but not biangulate, vein M, arising at about 
upper 1/4 of discocellulars, M not stalked with CuA r 

Male genitalia (Fig. 210, 243). Uncus possibly 
present as a small, triangular vestige at apex of tegumen. 
Saccus broad, but smaller than in Hydrelia. Labides 
about half length of tegumen, broadening and rounded 
at apices. Juxta narrow. Valva: narrow, sacculus with a 
weakly sclerotized, slender spine-like extension. 
Aedeagus: simple, vesica without cornuti. Sternum A8 

Female genitalia (Fig. 282). Ductus bursae long, 
weakly sclerotized throughout except for a short 
membranous break at middle; antrum absent. Bursa 
copulatrix: corpus bursae globose with dense covering 
of Eupithecia-Mke denticles (caeruleolineata) or minute 
denticles (pluristrigata) on anterior half; signum 
composed of fairly long radiating spines. 

Diagnosis. The male genitalia broadly resemble 
those of Hydrelia. Hastina, Macrohastina and 
Bihastina can be distinguished from other asthenine 
genera by the strongly dentate termen of both fore- and 
hindwings. Hastina differs from Macrohastina by the 
different wing colour and pattern and the absence of 
cornuti on the vesica, and from Bihastina by differ- 
ences in the venation and genitalia (see diagnosis 
under that genus). 

REMARKS. The posterior extension of the juxta 
takes the form of a narrow tongue in caerulineata, and 
it arises clearly from the juxta plate. In pluristrigata. 



however, the structure is more strongly sclerotized and 
appears to be derived from a union of sclerites at the 
base of the valvae. A tongue-like sclerite does occur, 
confusingly, in pluristrigata, but it is not apparently an 
extension of the juxta as it is a sclerotization of the 
dorsal diaphragmata (fultura superior). 

DISTRIBUTION. Burma, China, India, Japan, Russia. 

Species included. Three species. Sometimes 
caeruleolineata is treated as subspecies of subfalcaria. 
Genitalia examined: caeruleolineata (male, female), 
pluristrigata (male, female). 

Hastina caeruleolineata Moore, 1888 

Burma, China, India. 

Hastina pluristrigata (Moore, 1868) (Hyria?) 

China, India. 

Remarks. Although the shape of the hindwing of 
this species is almost the same as in caeruleolineata, 
the genitalia of both sexes are distinctive, the anal 
papillae being strongly sclerotized and forming a short, 
pointed ovipositor. It is further distinguished from 
caeruleolineata by the more weakly crenulated 
condition of the forewing termen. 

Hastina subfalcaria (Christoph, 1881) 


Japan, Russia. 

Macrohastina Inoue, 1982 

(Figs 144-146; 211, 244; 283.) 

Macrohastina Inoue, 1 982: 47 1 . Type species: Erosia 
azela Butler, 1878:403. 

MOTH (Fig. 144-146). Similar to Hastina in wing 
shape, but not colour and pattern. Head. Frons neither 
prominent nor broad. Labial palpi minute, not extend- 
ing beyond frons. Antenna ciliated. Wings. Colour and 
pattern distinctive, very similar in all three species. 
Venation as for Hastina except: forewing: areole single; 
hindwing with vein M arising from middle of 
discocellulars, and vein M stalked or not stalked with 
CuA r 

Male genitalia (Figs 211, 244). Saccus similar to 
that in Hastina or slightly shorter. Labides slightly less 
prominent than in Hastina. Juxta narrow. Valva: 
sacculus produced into a short spine or merely a pointed 
lobe. Aedeagus: vesica with cornuti in form of spines 
or denticles. 

FEMALE GENITALIA (Fig. 283). Ductus bursae broad 
posteriorly, narrowing toward middle of length; 
posterior half strongly sclerotized. Corpus bursae: 

signum composed of radiating denticles; large patch of 
prominent Eupithecia-style denticles also present in 
M. azela and M. gemmifera, minute denticles present 
in M. stenozona. 

Diagnosis. The three species have a distinctive and 
very similar wing colour and pattern making the genus 
easy to distinguish from all others. The sclerotizations 
of the corpus bursae are also characteristic. 

REMARKS. The three species in this genus are very 
similar in wing shape, colour and pattern and in genital 
structure. When describing the genus, Inoue (1982: 
471) noted that in M. azela and M. gemmifera vein M, 
of the hindwing is stalked with CuA r However, in 
M. stenozona M 3 is not stalked so the venational 
character should not be considered diagnostic for the 
genus. Nevertheless, we retain the genus Macrohastina 
because of other similarities between the three species, 
although it might be argued that this genus, Hastina 
and Bihastina should be treated as synonymous. 

Distribution. China, Japan, India, Nepal, Burma. 

SPECIES INCLUDED. Three species. Genitalia exam- 
ined: azela (male, female), gemmifera (male, female), 
stenozona (male, female). 

Macrohastina azela (Butler, 1878) (Erosia) 

Macrohastina gemmifera (Moore, 1868) 


India, Nepal, China. 

Macrohastina stenozona (Prout, 1926) 
comb. n. (Hastina) 

Hastina azela stenozona Prout, 1926. 
Hastina stenozona Prout; Xue & Zhu, 1999. 

Burma, India, China. 

Remarks. This species differs from the two others 
in the genus by the following characters: vein M, is not 
stalked with CuA ( on the hindwing; the sacculus of the 
valva bears a well-developed extension; comuti are 
arranged in two bundles of moderate length spines; the 
signum is much longer and narrower with denticles on 
the corpus bursae being very weak and small. How- 
ever, the wing pattern and colour are extremely similar 
to the other two species, and stenozona would appear 
therefore to belong to Macrohastina. 

Bihastina Prout, 1916 

(Figs 147-149; 212, 245; 284.) 

Bihastina Prout, 1916: 26. Type species: Bihastina 
albolucens Prout, 1916: 26. 


Moth (147-149). Head. Frons not protuberant. 
Labial palpi reduced, narrow, extending slightly beyond 
frons. Antenna not pectinate in male. Wings. Semi- 
translucent with extensive brown markings; termen of 
both wings deeply dentate. Forewing: two areoles 
present; vein R, arising before apex of second areole, 
not sharing stalk with IL .; vein R 5 arising from apex of 
second areole together with FL . Hindwing: Sc+R ( 
running close to costa for proximal half, almost touch- 
ing costa after the cell; discocellulars slightly curved, 
M, arising above middle of cell; vein M not stalked 

Male GENITALIA (Figs 212, 245). Uncus probably 
present in form of a broad, squat, membranous structure. 
Saccus broad. Labides small, weakly sclerotized, with 
setose apices. Transtillae well-sclerotized, uniting 
medially. Juxta with broad base and narrower exten- 
sion. Valva round terminally, without any extension or 
projection. Aedeagus: vesica lacking cornuti. Sternum 
A8 unmodified. 

Female genitalia (Fig. 284). Ductus bursae: 
antrum present in form of broad funnel; small 
colliculum present. Corpus bursae: signum appearing 
stellate, composed of long radiating spines. 

Diagnosis. The shape of the wings is similar to 
Hastina and Macrohastina. Bihastina differs from both 
these genera by the semi-translucent white wings 
marked with brown, the unstalked condition of vein R r 
the presence of two areoles on the forewing, and the 
rounded rather than elongated shape of the signum. 

Remarks. The structure at the apex of the tegumen 
is possibly the uncus, although it is membranous and 
not the typically narrow uncus occurring so widely in 
Geometridae. The labides are difficult to observe, but 
appear to be present, although small and inconspicu- 
ous, and situated just ventral to the well-sclerotized 
transtillae which are united to form a strong band 
dorsal to the aedeagus. 

Distribution. New Guinea. 

Species included. Three species. Genitalia exam- 
ined: albolucens (male, female), subviridata (male). 

Bihastina albolucens Prout, 1916 

West Irian. 

Bihastina subviridata (Bethune-Baker, 1915) 


Bihastina mera Prout, 1926. 
Papua New Guinea. 

Bihastina viridata (Warren, 1906) (Hastina) 
Papua New Guinea. 


Leucoctenorrhoe Warren, 1904 

(Figs 150; 213, 246; 285) 

Leucoctenorrhoe Warren, 1904: 526. Type species: 
Leucoctenorrhoe quadrilinea Warren, 1904: 527. 

Moth (Fig. 150). Head. Frons protuberant as in 
Hydrelia. Labial palpi extremely narrow and short. 
Antenna in both sexes bipectinate, pectinations very 
long. Wings. White; lines less wavy than in usual 
Asthenini condition ; forewing somewhat narrower and 
rounder than in most Asthenini. Forewing: areole single, 
vein R j diverging from common stem distal to point at 
which R ? diverges; vein M 2 arising slightly closer to M 
than to M 3 in both fore- and hindwing. Hindwing 
slightly angled at middle of termen; discocellulars 
straight, not angled; vein M, stalked with Rs; M 
stalked with CuA ]5 length of stalk variable. 

Male genitalia (Fig. 213, 246). Labides with broad 
apices. Juxta flask-shaped. Valva short, broad, rounded, 
lacking any extension; apex fringed with hair-like 
scales expanded at their tips, each scale about as long 
as valva. Aedeagus short and broad; vesica with small 
sclerotization, lacking spine-like cornuti. Sternum A8 

Female genitalia (Fig. 285). Anal papillae short 
and broad; apophyses strong. Ductus bursae short, 
with small colliculum. Corpus bursae globose; signum 
pear-shaped, composed of radiating denticles. 

DIAGNOSIS. Leucoctenorrhoe quadrilinea is distin- 
guished from Asthena by the presence of bipectinate 
antennae in both sexes and by the different venation. 
Leucoctenorrhoe differs from those other genera in 
which vein R t diverges distal to R 5 on the forewing, 
and in which hindwing vein M, is stalked, by the 
bipectinate antenna, the long cell on the hindwing and 
the pure white ground colour of both wings. The male 
genitalia differ from all other Asthenini notably in the 
short valva with the presence of long scale-like hairs 
fringing its apex and the short, broad aedeagus. 
Although the hairs around the apex of the valva occur 
also in Epicyme, the antennae, venation, aedeagus and 
female genitalia are different. See diagnosis of Epicyme. 

Remarks. The Peruvian Leucoctenorrhoe 
quadrilinea is the only species of Asthenini recorded 
from the Neotropics. 

Distribution. Peru. 

SPECIES INCLUDED. Monotypic. Genitalia examined: 
quadrilinea (male, female) 

Leucoctenorrhoe quadrilinea Warren, 1904 



Parasthena Warren, 1902 

(Figs 151; 214, 247; 260, 261; 286.) 

Parasthena Warren, 1902: 361. Type species: 
Parasthena flexilinea Warren, 1902: 362;Holloway, 
1997: 183. 

MOTH (Fig. 151). Head. Frons less broad and promi- 
nent than in Hydrelia. Antenna simple, nearly smooth 
in both sexes. Labial palpi minute. Wings. Pale grey- 
brown with darker multiple fasciae and black discal 
spots. Forewing shaped as in Hydrelia, termen weakly 
angled; male with small anal lobe; cell short and broad; 
areole single; vein R 5 separate, arising from below 
angle of areole. Hindwing: male with posterior margin 
folded under wing; termen slightly produced at M and 
ML; cell short and broad, Sc+R ( united with Rs to 3/4 of 
cell, M, stalked with Rs, discocellulars not biangulate. 

Male genitalia (Figs 214, 247; 260, 261; 
286). Tegumen long and narrow. Saccus short and 
narrow. Labides short. Juxta flask-shaped. Valva broad, 
covered with long hairs; sacculus without a projection. 
Aedeagus: vesica lacking cornuti. Sternum A8 with 
anterior margin produced into pair of short but con- 
spicuous apodemes; tergum A8 with anterior margin 
produced into a single conspicuous, medial projection. 

Female genitalia (Fig. 286). Ductus bursae: 
antrum leading into short membranous section of duc- 
tus; colliculum long and conspicuous. Corpus bursae: 
oval rather than globose, with scattering of very weak 
denticles in region of signum; signum prominent, 
elongated, composed of long radiating spines. 

Diagnosis. The genitalia exhibit many similarities 
to those of Poecilasthena: the tegumen is similarly 
extended and the saccus is short and of a similar shape; 
the juxta is of the same form; sternum A8 in the male is 
modified in both genera; and, in the female, the 
narrowness of the ductus bursae with its associated 
colliculum are similar. Parasthena is, however, 
distinguished from Poecilasthena by having a single 
areole, an unexpanded costa of the valva, and a 
prominent colliculum. 

Remarks. Holloway (1997: 183) removed 
Parasthena from synonymy with Hydrelia, noting that 
it lacked certain characters of the Asthenini. It is very 
close to Poecilasthena, but we have not synonymized 
the genera because of the presence of a single areole 
and a much larger colliculum in Parasthena. 

We have also examined further specimens from 
Seram and Papua New Guinea of material representing 
what was noted by Holloway (1997: 184) as 'a related, 
somewhat more strongly marked, undescribed species' . 
In the forewing, the costa is heavily marked with dark 
grey, especially toward apex; the termen is more exca- 
vated between the apex and vein M than in flexilinea; 


and the angle of the wing is more pronounced. In the 
hindwing, the termen is more strongly dentate than in 
flexilinea, notably at M and M v The median band on 
the forewing and the postmedian band on both wings 
are darker than in flexilinea. The genitalia, however, 
are not convincingly different, and we are undecided 
about the taxonomic status of this material. 

Distribution. Sulawesi; Philippines; Borneo; 
Seram; Papua New Guinea. 

Species included. A single named species. Genita- 
lia examined: flexilinea (male, female). 

Parasthena flexilinea Warren, 1902 

Sulawesi, Philippines, Borneo. 

Poecilasthena Warren, 1894 

(Figs 152-171; 215-218; 262-265; 287-290, 295) 

Poecilasthena Warren, 1894: 394. Type species: 
Acidalia pulchraria Doubleday, 1843: 286. 

Astheniodes Hampson, 1903: 647. Type species: 
Astheniodes polycymaria Hampson, 1903a: 648. 

Poecilasthena Warren; Holloway, 1997: 182. 

Moth (Figs 152-171). Head. Frons narrow, not 
prominent. Labial palpi minute, extended slightly 
beyond frons. Antenna simple, with short cilia. Wings. 
Ground colour of fore- and hindwings generally white, 
marked with numerous transverse lines, pale grey in 
papuensis group; lines usually pale green to greyish 
green; wings brown in some species. Forewing: 
triangular; cell with a minute spot; termen slightly 
curved, anal angle well expressed; cell longer than 
half-length of wing; areole double; vein R t arising 
from before apex of areole; R,. arising from apex of 
areole together with R Hindwing: termen slightly 
angled medially, sometimes rounded, anal margin 
slightly longer than costa, discocellulars strongly 
biangulate or not biangulate (papuensis group). 

Male genitalia (Figs 215-218, 262-265). Saccus 
ranging from broad to narrow. Labides curved, form- 
ing a harp-like structure, extending as posteriorly 
directed, broader membranous projections from base 
of costa. Juxta flask-shaped. Valva covered with setae; 
sacculus not extended; costa broadened at middle, 
setae absent from broadened part of costa; coremata 
prominent, arising from base of valva, or absent. 
Aedeagus short; vesica with cornuti arranged in a 
bundle of short spines, sometimes in two bundles, or 
cornuti absent. Sternum A8 often with proximal margin 
produced on each side so appearing W-shaped, some- 
times the two extensions long and pointed. 

Female genitalia (Figs 287-290; 295). Ductus 
bursae short, with small colliculum at lower part. 
Corpus bursae usually globose or subglobose, rarely 



more elongated; signum round or oval, composed of 
spines radiating from denticulate core, sometimes with 
marked medial ridge (as in Fig. 295). 

Diagnosis. In most species of Poecilasthena the 
discocellulars of the hindwing are strongly biangulate, 
unlike the situation m Asthena. The two species in the 
papuensis group, where the discocellulars are not 
biangulate, are distinguished from Asthena mainly by 
differences in the genitalia. In the male, the costa of the 
valva is broadened and hairless, in many species there 
are a pair of well developed coremata attached to the 
base of the valva, and sternum A8 often bears a pair of 
projections from the anterior margin. In the female, the 
ductus bursae is membranous other than for a small 
colliculum, and the signum is often circular, with the 
spines in the middle being minute and those extending 
from the margin being very long. Poecilasthena exhibits 
many similarities to Parasthena but has a double areole 
(single in Parasthena). 

REMARKS. Some variation occurs within the genus. 
In pulchraria and subpurpureata (Walker) the saccus 
is smaller and the sacculus very short with a small 
needle-like terminal process arising from the base of 
the valva and separating from it slightly. Coremata are 
absent and the cornuti are arranged in two bundles. 
Sternum A8 is weakly modified with both ends of the 
proximal edge extended proximally. 

In papuensis and euthecta, the discocellulars in the 
hindwing are not biangulate and tergum A8 of the male 
has a medial projection extending from the anterior 
margin. In the female, the antrum is larger and the 
signum is smaller with the spines at the middle larger 
and gradually becoming longer toward the edges. 

Notable similarities exist between Poecilasthena 
and the Eupitheciini in the form of the labides in the 
male genitalia and the colliculum in the female genita- 
lia. In Eupithecia the juxta is shaped characteristically 
like an hourglass. The 'lower' (anterior) part of each 
labides takes the form of a narrow, elbowed rod, one 
end of which extends into the waist of the hourglass. 
The 'upper' (posterior) part of the labides, which is less 
well sclerotized, is broader than the lower section. In 
Poecilasthena the juxta is flask-shaped with a broad 
anterior element from which extends a narrow, rod- 
like component. Although the shape of the juxta differs 
from that present in Eupithecia, the labides are notably 
similar. The anterior sclerite narrows and almost meets 
the rod-like component of the juxta. The posterior part 
is broader and more membranous. In the female, the 
ductus bursae is narrow in Poecilasthena and 
Eupithecia (and in Parasthena) and there is a small 

Distribution. Australia, Burma, Fiji, Indonesia, 
Malaysia, New Caledonia, New Zealand, Papua New 
Guinea, Philippines. 

Biology. Details of life histories were recorded by 
McFarland (1979: 41 ,42) for two species (P. pulchraria 
and an undetermined species) and, for P. pulchraria by 
McFarland, 1988: 352-354), and were summarized by 
Holloway ( 1 997: 182). The larva fed on leaves, flowers 
and fruits of Astroloma humifusum (Epacridaceae, the 
Australian Heaths), a plant considered likely to be the 
wild host. The second species (not 'lischnophrica 
Turner' as suggested by McFarland, 1979, see 
Holloway, 1997), was found feeding on Leptospermum 

SPECIES INCLUDED. 32 species. Genitalia examined: 
aedoea (male), anthodes (male), burmensis (male), 
character (male, female), dimorpha (male, female), 
leucydra (male, female), nubivaga (male, female), 
papuensis (male, female), prouti (female), pulchraria 
(male, female), scoliota (female), schistaria (male, 
female), subpurpureata (male, female), xylocyma 
(male, female). 

Poecilasthena aedoea Turner, 1926 


Poecilasthena balioloma (Turner, 1907) 



Poecilasthena burmensis Prout, 1926 


Poecilasthena character Prout, 1932 

Indonesia, Malaysia. 

Poecilasthena cisseres Turner, 1933 


Poecilasthena dimorpha Holloway, 1979 

New Caledonia. 

Remarks. This species exhibits features differing 
from typical Poecilasthena. In the forewing, vein R 5 
arises from below the apex of the areole, whereas in 
other species of this genus it typically arises at the 
apex. The posterior margin of the hindwing is longer 
than in other species belonging to the genus and the 
termen is rounded and not angled. In the male, the 
valva is distinctive being broad with a narrow base and 
a dense arrangement of long hairs fringing its distal 
margin. In the female, the corpus bursae is elongated 
rather than globose or subglobose and the signum is 
elongated, not rounded. 

Poecilasthena euphylla (Meyrick, 1891) 



Poecilasthena fragilis T\irner, 1942 


Poecilasthena glaucosa (Lucas, 1888) (as 


Euchloris (lodis) microgyria Lower, 1 894 

Poecilasthena inhaesa Prout, 1934 


Poecilasthena iopolia (Turner, 1926) 



Poecilasthena ischnophrica Turner, 1941 


Poecilasthena leucydra Prout, 1934 

Fiji, New Caledonia. 

Poecilasthena limnaea Prout, 1926 

New Guinea. 

Poecilasthena nubivaga Prout, 1932 

Indonesia (western). 

Poecilasthena oceanias (Meyrick, 1891) 



Poecilasthena panapala Turner, 1922 


Poecilasthena paucilinea Warren, 1906 

Papua New Guinea. 

Poecilasthena phaeodryas Turner, 1931 


Poecilasthena pisicolor Turner, 1942 


Poecilasthena prouti West, 1929 (as 



Poecilasthena pulchraria (Doubleday, 1843) 



Asthena ondinata Guenee, [1858] 
Chlorochroma plurilineata Walker, 1861 
Asthena pulchraria decolor Turner, 1904 

New Zealand, Australia. 

FOODPLANTS. Epacridaceae: Astroloma; Brachyloma; 
Epacris; Monotoca. 

Poecilasthena scoliota (Meyrick, 1891) 



Poecilasthena sthenommata Turner, 1922 


Poecilasthena suhpurpureata (Walker, 

[1863]) (Asthena) 

Astheniodes polycymaria Hampson, 1903 
Acidalia tuluiata Felder & Rogenhofer, 1875 

New Zealand. 

REMARKS. The holotype of polycymaria is labelled 
as being from India, which, if correct, would give a 
very peculiar distribution pattern for suhpurpureata. 
Comparison of the genitalia of the male holotype of 
polycymaria with those of a specimen of suhpurpureata 
from New Zealand shows a remarkable good match 
between the structures and there seems to be no reason 
to treat polycymaria as a separate species. Prout (1927: 
76) regarded the type locality as being 'evidently in 
error', and it is difficult to dispute that comment. 

Poecilasthena thalassias (Meyrick, 1891) 


Asthena pellucida Lucas, 1892 
Australia, Papua New Guinea. 

Poecilasthena urarcha (Meyrick, 1891) 


Australia (including Tasmania). 

REMARKS. The wings of this species are green and 
the termen of the hindwing shows no sign of an angle. 

The papuensis group 

Two species previously assigned to Asthena are here 
transferred to Poecilasthena. One of them was origi- 
nally described under Hydrelia. Their association with 
Asthena is explained by the fact that, unlike the con- 
dition in other Poecilasthena, the discocellulars in the 
hindwing are not biangulate. However, the genitalia 
closely resemble those of other Poecilasthena and 
sternum A8 is modified similarly to Poecilasthena. See 
also 'Remarks' under the generic description above. 



Poecilasthena euthecta (Turner, 1904) 
comb. n. (Asthena) 


Remarks. Nielsen, Edwards & Rangsi (1996) treated 
this species in Minoa. However, the genitalia structures 
are not consistent with those of Minoa. 

Poecilasthena papuensis (Warren, 1906) 
comb. n. (Hydrelia) 

Papua New Guinea. 

The anthodes group 

The moths of this cohesive species group are brown. 
The labides are fairly long, nearly extending to the end 
of the tegumen, the saccus is narrow, coremata extend 
from the base of each valva and the vesica lacks 
cornuti. Sternum A8 is markedly elongate being twice 
the length of the other sterna; the proximal end is 
rounded, and from 1/3 toward the distal end it become 
very narrow, before broadening a little distally. Tergum 
A8 is not modified. The female genitalia are indistin- 
guishable from those of other Poecilasthena species. 

Poecilasthena anthodes (Meyrick, 1891) 
comb. n. (Asthena) 


Poecilasthena schistaria (Walker, 1861) 
comb. n. (Acidalia) 

New Zealand. 

Poecilasthena xylocyma (Meyrick, 1891) 
comb. n. (Asthena) 


Polynesia Swinhoe, 1892 

(Figs 172-174; 219, 252; 291.) 

Polynesia Swinhoe, 1892: 4. Type species: Pomasia 
sunandavaWalker, 1861: 657; Holloway, 1997: 189. 

Placotome Warren, 1894: 395. Type species: Poly- 
nesia truncapex Swinhoe, 1892: 5. 

Moth (Figs 172-174). Head. Frons narrow, not pro- 
tuberant. Labial palpi long and robust, not slender, 
extending about 1/4 to 1/3 beyond front of head. 
Antenna in both sexes simple, with very short cilia. 
Legs. Hind tibia of male with terminal spurs only; hind 
femur of male expanded and with thick hair pencil in 
groove. Hind tibia of female with all spurs present. 
Wings. Primrose yellow variously marked with irregu- 
lar, rufous spots. Forewing with costa somewhat 
broadened near base, termen very weakly crenulated, 
slightly produced medially, concave under CuA,; areole 

single, small, all radial veins stalked, R t diverging 
from stalk distal to divergence point of R 5 ; male 
forewing of truncapex truncated so costa appears 
angled; tip of angle folded over dorsal surface. 
Hindwing: apex rounded, termen produced both at M, 
and CuA,; cell short, not longer than 1/3 length of 
hindwing; discocellulars gently curved; M 2 arising a 
little above middle of discocellulars, Rs and M,, M 3 
and CuAj stalked; sometimes underside of male 
hindwing with hair tuft at base of cell or hair ridge 
along wing. Abdomen. Long and narrow in both sexes; 
tympanal organ with length exceeding that of first 
visible sternum (A2); anterior angles of second visible 
sternum (A3) also produced. 

Male genitalia (Figs 219, 252). Saccus quadrate. 
Tegumen narrow and long, uncus vestigial. Labides 
prominent, forming long, well-sclerotized, curved 
spines. Juxta in form of fairly small, irregular plate. 
Valva fairly broad, simple, setose; sacculus not 
projecting beyond margin of valva, elbowed termi- 
nally, but without terminal projection. Aedeagus simple, 
cornuti lacking. Last tergum and sternum elongated, 
sternum usually longer than tergum. 

Female genitalia (Fig. 291). Ductus bursae 
membranous posteriorly, with long antrum anteriorly. 
Corpus bursae: globose; signum composed of radiat- 
ing denticles; small additional signum present near 
cervix bursae in sunandava. 

Diagnosis. This genus is easily distinguished from 
other Asthenini by the primrose yellow wing colour 
dotted with irregular rufous spots and the venation. 
The smooth, large hook-like labides in the male 
genitalia are highly distinctive. 

The wing colour of Polynesia is very similar to that 
of Pseudopolynesia Holloway. Polynesia is easily 
distinguished from Pseudopolynesia in the structure of 
the genitalia. The male of Pseudopolynesia lacks the 
conspicuous labides of Polynesia and the shape of the 
tegumen differs between them. In the female of 
Pseudopolynesia there is no signum and the bursa 
copulatrix is double. 

Distribution. India, China, Thailand, N.E. 
Himalaya, Malaysia, Indonesia, Sri Lanka, Papua New 

Remarks. The labial palpi are longer than is usual in 
Asthenini and certain features of the male genitalia are 
peculiar. However, typical asthenine characters are the 
reduced uncus, the signum being composed of radiat- 
ing denticles, and the presence of a small areole on the 

Species included. Three species. Genitalia exam- 
ined: curtitibia (male, female), sunandava (male, 
female), truncapex (male, female). 

Polynesia curtitibia Prout, 1922 

India, Thailand, N.E. Himalaya, Malaysia. 

Polynesia sunandava (Walker, 1861) 


Cambogia? aeriferata Walker, [1863]. 

Sri Lanka, India, Malaysia, Indonesia, China, Papua 
New Guinea. 

Polynesia truncapex Swinhoe, 1892 

India, China, Malaysia, Indonesia. 

Anydrelia Prout, 1938 

(Figs 175-177; 220,253; 292.) 

Anydrelia Prout, 1938: 177 (see Prout, 1934-1939). 
Type species: Brabira plicataria Leech, 1897: 72. 

MOTH (Figs 175-177). Head. Frons broad, promi- 
nent in male, flat in female. Labial palpi minute, not 
extended beyond frons. Antenna: weakly serrate in 
male with short cilia, simple in female. Wings. Brown, 
with darker but weak postmedial band and even weaker 
subterminal line: hindwing much smaller than forewing 
in both sexes. Forewing broad, costa much longer than 
dorsum, apex slightly produced, termen rounded; areole 
single; vein R : and R, _ 5 diverging beyond (distal to) 
areole; vein R s diverging almost at end of areole and 
proximal to divergence point of R^ vein M, diverging 
from areole, discocellulars curved, M, arising slightly 
closer to M, than to M r Hindwing of both sexes 
markedly smaller than forewing, termen rounded, anal 
lobe in male expanded and folded under wing, ventral 
surface with numerous specialized scales, dorsal 
surface with long hair-scales; Sc+R L combined with 
Rs to 3/4 of cell, which is as long as half length of the 
wing; Rs united for short distance with M p M ( curved 
downwards, discocellulars strongly biangulate, M, 
arising from the second angle, very near M . 

Male genitalia (Figs 220, 253). Saccus short. 
Labides reduced to pair of setose membranous heads. 
Juxta broad basally, narrowing slightly towards apex, 
which is divided. Valva broad, with the costa incurved, 
sacculus not sclerotized, but with a small sharp terminal 
process arising from just beyond the middle valva. 
Aedeagus: vesica lacking cornuti. Sternum A8 

Female genitalia (Fig. 292). Ductus bursae weakly 
sclerotized throughout. Corpus bursae covered with 
minute denticles; signum absent. 

Diagnosis. This genus is very easy to distinguish 
from other asthenine genera by the specialized 
hindwing in the male and by the absence of the asthenine 
signum in the female. Typical Asthenini characteris- 


tics include a broad, protuberant frons, minute labial 
palpi, a reduced uncus, short labides and a sclerotized 
ductus bursae. 

Distribution. China, India, Nepal. 

Remarks. The absence of a typical asthenine signum 
is interpreted as a loss. 

Species included. Three species have been 
described in this genus. Genitalia examined: distorta 
(male, female), plicataria (male, female). 

Anydrelia dharmsalae (Butler, 1883) (Ephyra) 

Anydrelia distorta (Hampson, 1895) 

India, Nepal, China. 

Anydrelia plicataria (Leech, 1897) (Brabira) 


The following genera exhibit some similarity to the 
Asthenini, but we have excluded them from the core 
group for reasons given under each genus. Three of the 
genera included below are treated in the same detail as 
those above: Minoa Treitschke, which is one of the 
three genera included by Pierce ( 1 9 1 4) in his definition 
of the Asthenini; Cleptocosmia Warren, which was 
listed by its describer in the original use of the family 
group name Astheninae; and Chalyboclydon Warren, 
the composition of which is complex (see above). 

Minoa Treitschke, 1825 

(Figs 178. 179; 221. 254; 267; 293.) 

Minoa Treitschke, 1825: 445. Type species: Geometra 
euphorbiata [Denis & Schiffermuller], 1775: 1 16 (a 
junior subjective synonym of Phalaena murinata 
Scopoli, 1763:229). 

Moth (Figs 178, 179). Head. Frons not prominent. 
Labial palpi more rough scaled than in most Asthenini 
and not strongly reduced. Antenna: male with short 
cilia. Wings: drab, dark to medium grey-brown or 
ochreous, margins smooth, not crenulated; costa of 
forewing short, termen relatively straight. Forewing: 
areole double; vein R : arises from well before apex of 
second areole, veins R„ R and R 5 arise, independ- 
ently, from its apex. Hindwing: discocellulars oblique, 
not biangulate. 



Male genitalia (Figs 221, 254; 267). Saccus 
rounded. Labides digitate, curved. Juxta in form of a 
large plate produced posteriorly into a long, finger-like 
process. Valva distinctive, narrow; costa and sacculus 
strongly sclerotized each terminating as a spine. 
Aedeagus: broad; vesica with a single, long cornutus or 
a single short cornutus subtended by a sclerite. Abdo- 
men (Fig. 267) terga and sterna well-sclerotized in both 
male and female; sternum A8 trapezoidal in male. 

Female genitalia (Fig. 293). Anal papillae short, 
membranous. Bursa copulatrix: ductus bursae promi- 
nent, broad, strongly and uniformly sclerotized, 
extending into small corpus bursae, which is suddenly 
demarcated from remaining membranous corpus; 
signum, when present, small, composed of a few spines, 
not of radiating denticles and spines. 

DIAGNOSIS. This distinctive genus is recognizable 
particularly by the uniform wing colour of the moths, 
the shape of the valva, the form of the ductus bursae, 
and the strongly sclerotized condition of the sterna and 
terga of the abdomen in both sexes. 

Distribution. Europe, including Eastern Europe. 

Remarks. Minoa was included in 'Astheninae' by 
Pierce (1914: 38) in the paper that first defined the 
group, but there is no sound reason to associate it with 
the core of the tribe. Although the uncus is reduced and 
labides are present, the labial palpi are relatively robust, 
and while the sacculus is extended, the extension does 
not resemble that typical in Asthenini, indeed, the 
overall shape of the valva is unusual. The female 
genitalia are highly distinctive and lack the typical 
asthenine signum. The tribal association of Minoa 
remains unclear. 

Minoa is considered currently to include just one 
polytypic species. However, a preliminary study of the 
genitalia of specimens from several localities suggests 
that there are, in fact, two species. For the purposes of 
the present paper we accept the identity of the type 
species as the widespread species, although this re- 
quires confirmation. Further study is being undertaken 
to establish the content of the genus and the names that 
should be associated with the species involved. 

Species included. Probably two species, one of 
which is described. Genitalia examined: murinata 
murinata (male, female), murinata amylaria (male, 

The variation within Minoa needs re-examination. 
There appear to be two species rather than one, although 
currently one is accepted (Karsholt & Razowski, 1996: 
248). One of these species is widespread. Specimens 
of the other in the collection of the BMNH are from 
Sicily and Dalmatia. It is likely from an examination of 
the original description that the widespread species is 
what is generally perceived to be murinata, but this 

question deserves further study. We have yet to dis- 
cover if an available name exists for the second species. 

Minoa murinata murinata (Scopoli, 1763) 


Minoa cyparissaria Mann, 1 854. 

Geometra euphorbiata [Denis & Schiffermiiller], 1775. 

Phalaena fuscata Hufnagel, 1767. 

Acidalia italicata Milliere, 1885. 

Phalaena (Geometra) sordiata Linnaeus, 1767. 

Ph[alaena] Geomfetra] unicolorata Hiibner, [1787]. 

Widespread in Europe, Russia, Asia Minor, Central 

FOODPLANTS. Euphorbiaceae: Euphorbia amygda- 

Minoa murinata amylaria Prout, 1914 

Alps, Italy. 

Minoa murinata limburgia Lempke, 1969 


Minoa murinata lutea Schwingenschuss, 


Chalyboclydon Warren, 1893 

(Figs 180; 222, 256; 294.) 

Chalyboclydon Warren, 1893: 366. Type species: 
Chalyboclydon ma rginata Warren, 1893: 366. 

Description of Chalyboclydon marginata 

MOTH (Figs 180). Head. Frons neither broad nor 
protuberant. Labial palpi slender, reduced, but extend- 
ing further from head than in most Asthenini genera. 
Antenna simple in both sexes, ciliated. Wings. Broad, 
off-white with dark markings at termen similar to 
those in Eschatarchia lineata. Forewing broad, apex 
slightly produced, termen angled medially, weakly 
crenulated above angle, straight below angle; hindwing 
with termen angle more marked than in forewing, 
produced as in E. marginata; cell of both wings 
extending well beyond 1/3 length of wing, 
discocellulars strongly oblique; forewing with small 
single areole, R { stalked, R^ diverging from stem after 
R 5 and well beyond end of areole; hindwing with 
Sc+R, combined with Rs for 3/4 length of cell, Rs and 
M { stalked, M 2 arising from above middle of 
discocellulars, M 3 not stalked. 

Male genitalia (Figs 222, 256). Saccus broad. 
Labides narrow, weakly sclerotized. Juxta narrow. Valva 
narrow; costa weakly sclerotized, gently convex 



medially and with a denticle arising subterminally; 
sacculus well sclerotized, with broad lobe medially. 
Aedeagus: vesica with group of cornuti. Abdominal 
sternum A8 unmodified. 

Female genitalia (Fig. 294). Ductus bursae short 
and sclerotized merging with long corpus bursae. 
Corpus bursae with one signum in form of a prominent 
band from which extends strong spines; second signum 
irregular, bearing a number of strong denticles; neither 
signum composed of radiating denticles or spines. 
Sternum A7 forming a well-sclerotized, crescent- 
shaped plate near ostium bursae. 

DIAGNOSIS. The wing colour, pattern and venation 
resemble, to some degree, those features in 
Eschatarchia. In both, the angle in the middle of the 
hindwing termen is pointed. Chalyboclydon marginata 
may be distinguished by its much longer palpi and the 
very weakly angled forewing termen. The genitalia 
differ significantly (compare figures). The wing pattern 
distinguishes C. marginata from other genera. 

Distribution. India, Sikkim, Burma, China. 

REMARKS. The genus lacks the typical asthenine 
signum, and the valvae are narrower and their 
sclerotizations more complex than in the Asthenini. 

Chalyboclydon was described by Warren (1893: 
366) as a monotypic genus, from two localities 'Sikkim' 
and 'Momeit, Burmah', in a paper including descrip- 
tions of various genera and species in the H.J. Elwes 
collection, housed in the BMNH. Although Warren's 
description was said to be of the female sex, all 
three syntypes of marginata are males. There is a 
further specimen, a female, labelled as being from 

The species C. flexilinea was described later (War- 
ren, 1898: 22) from 'One specimen from the Khasis. 
Closely allied to but apparently distinct from the Sikkim 
marginata! This specimen, the holotype, is a male and 
bears a label, in Warren's hand, 'Chalyboclydon 
flexilinea Warr Type $'. A second label, by Prout, 
states: 'Chalyboclydon marginata Warr.' Warren gave 
a figure of 36 mm as the wingspan of the species, 
which is over 10 mm more than actually is the case for 
the syntypes of marginata and also is much greater 
than the measurement of flexilinea. While Warren's 
description of marginata might fit both species, the 
colour plate (23: 16) is clearly of marginata. Thus 
despite the confusing label of Prout, and the anoma- 
lous wing measurement given by Warren, the identities 
of the two species do not seem to be in doubt. However, 
to avoid further confusion, we have fixed the identity 
of Chalyboclydon marginata Warren, see below. 

Species included. One species known. Genitalia 
examined: marginata (male, female). 

Chalyboclydon marginata Warren, 1893 

Hydrelia? apicata Wileman, 1916. 

REMARKS. To fix the identity of marginata, the male 
syntype housed in the BMNH, illustrated in Fig. 180, 
and bearing the following label data is here designated 
as LECTOTYPE: Lectotype; Sikkim. Moller. 1888; 
Chalyboclydon marginata Warr. Type B&[handwritten 
by Warren]; Collectio[n] H.J. Elwes; Rothschild 
Bequest B.M. 1939-1. 

Distribution. India, Sikkim, Burma, China. 

Cleptocosmia Warren, 1896 

(Figs 181:223,256.) 

Cleptocosmia Warren, 1896: 383. Type species: 
Cleptocosmia mutabilis Warren, 1896: 383. 

Moth (Fig. 181). Head. Male. Frons flat, not broad. 
Labial palpi broad and long, projecting well beyond 
head, with segment 3 strongly elongated. Antenna 
simple, ciliated. Hindtibia bearing many hair-like 
scales, with a very long hair tuft at base, the two outer 
spurs of hind tibia also with hair scales. Wings. Orange 
brown; similar in shape to those of Chalyboclydon 
marginata but with forewing slightly narrower and 
apex rather more sharply angled; termen weakly angled 
medially; hindwing weakly angled medially. 
Forewing with hyaline fovea, and with the basal half of 
wing clothed with erect hair-like scales creating a fur- 
like appearance. Venation: cell very short, no longer 
than 1/3 length of forewing or hindwing; discocellulars 
nearly straight on forewing, angled and oblique on 
hindwing; forewing with areole absent, radials stalked 
with vein R 5 diverging from common stem before 
(proximal to) divergence point of R : ; R, _. arising before 
upper angle of cell, M not stalked, M, arising slightly 
closer to M t than to M,; hindwing with Sc+R combined 
with Rs to 3/4 of cell, Rs stalked with M |( M, stalked 
with CuA p CuA 2 arising near lower angle of cell. 
Female unknown. 

Male genitalia (Fig. 223, 256). Saccus rounded. 
Labides absent. Juxta broad, weakly sclerotized. Valva: 
broad, setose; costa with short point extending at 
middle; sacculus not extended, folded near base of 
valva and strongly setose. Aedeagus: broad; vesica 
with two large spines and some small ones. 

DIAGNOSIS. Cleptocosmia mutabilis may be recog- 
nized by the fur-like appearance of the basal half of 
forewing in the male. The erect, sharp angle at the 
middle of the costa of the valva, and the presence of 
two huge spines on the vesica of the aedeagus is also 
very characteristic. The venation is almost the same as 
that of 'Chalyboclydon' flexilinea, but, besides the fur- 
like scaling on the forewing, it can also be distinguish 
from that genus by long palpi, very weakly marked 
wings and the form of the male genitalia. 


Distribution. Australia. 

Remarks. Although Warren originally placed this 
genus in the Astheninae' (i.e., Asthenini), Nielsen, 
Edwards & Rangsi ( 1 996) excluded it from the tribe by 
listing it under a heading 'Unplaced to tribe' . The long 
labial palpi, the normal frons, the hairy hind tibia and 
forewing are all unusual. The female is unknown, so 
we are unable to comment on the form of the signum. 
On balance, however, like Nielsen etal. (1996) we are 
unable to place Cleptocosmia to tribe. 

The species is know from only the male holotype in 
the BMNH, and a further male recently discovered in 
the accessions of the ANIC. We are grateful to Mr E.D. 
Edwards for searching the ANIC collections. 

Species included. One species. Genitalia exam- 
ined: mutabilis (male). 

Cleptocosmia mutabilis Warren, 1896 


Eois Hiibner, 1818 

(Figs 182-184, 224, 257.) 

Eois Hiibner, 1818: 27. Type species: Eois russearia 

Hiibner, 1818:27. 
Cretheis Meyrick, 1886: 192, type species Cretheis 

cymatodes Meyrick, 1886: 193. 

In its broadest sense (e.g., Holloway, 1997, and as 
catalogued in Scoble, 1999), Eois is a large genus with 
c. 250 species, the type species of which was described 
from Surinam. A list of the generic synonyms is also 
provided by Holloway (1997: 184), who examined the 
type species of the names involved. Holloway sug- 
gested that despite some doubts that the Old World 
species of Eois might not be congeneric with those 
from the New World, most species were correctly 
united under the single genus. The basis of Holloway 's 
argument lies chiefly in the presence of a multispined 
signum set into the wall of the corpus bursae. Eois is 
certainly in need of a species-level review, an exercise 
that would help further investigation of the question of 
its monophyly. 

If the suggestion is accepted that Eois is mono- 
phyletic, with relatively minor adjustment to its 
composition, the question arises over its position in 
relation to the Asthenini. Notable asthenine characters 
include small labial palpi and a strongly reduced/lost 
uncus. The shape of the valva is also remarkably 
similar to that in Asthenini. However, unlike the posi- 
tion in Asthenini, labides are absent and the signum is 
distinctive, differing markedly from that seen in typi- 
cal Asthenini . Even if the Old World species should be 
shown not to be congeneric with those of the New 
World, we would exclude them from the Asthenini 
because of these features. 


Cretheis (Figs 1 82, 224, 257) is probably a synonym 
of Eois Hiibner although the Australian species of Eois 
were included in Cretheis by McQuillan & Edwards 
(in Nielsen, Edwards and Rangsi, 1996: 228). As with 
many species of Eois it shares some features of 
Asthenini, including short, narrow palpi, a strongly 
reduced uncus, and the presence of a small extension 
of the sacculus. It was included in the tribe by 
McQuillan & Edwards (in Nielsen, Edwards & Rangsi, 
1996: 227), but is excluded here because of the very 
different signum in the female, which is robust, 
multispined and has its base usually set in an evagina- 
tion of the bursa wall. Furthermore, the frons is very 

Species included. Two hundred and forty seven 
species of Eois (including those in Cretheis) are listed 
in Scoble (1999). We have not reviewed the species of 
this large genus, so they are not listed here. 

Pseudopolynesia Holloway, 1997 

(Figs 185,225.) 

Pseudopolynesia Holloway, 1997: 190. Type-species: 
Pomasia amplificata Walker, 1 86 1 : 658. 

One described species was included in the genus by 
Holloway (1997), who pointed out that there were at 
least two in the group, which extends from Sundaland 
to New Guinea. The remarkable similarity in the colour 
pattern of Pseudopolynesia and Polynesia encouraged 
us to study the genus to see if there was an asthenine 
association of Pseudopolynesia. 

The labial palpi are prominent and extended strongly 
in front of the head. In the male genitalia (Fig. 225), the 
tegumen is dome-shaped and the uncus appears to be 
absent. The sclerotizations of the anellus are complex, 
but labides do not seem to be present. In the female, the 
signum is absent and the corpus bursae is double with 
a posterior, spinose component and a flimsy anterior 

Characters that support the exclusion of 
Pseudopolynesia from the Asthenini are the presence 
of well-developed labial palpi, the absence of labides, 
the lack of an extension of the sacculus and the absence 
of an asthenine signum. While an uncus is absent from 
Asthenini and Pseudopolynesia, this situation is not 
unique to the tribe and does not define it alone. Denticles 
are present on the bursa copulatrix of Pseudopolynesia 
and Eupithecia, although those in Pseudopolynesia are 
not as robust. 

Despite the similarity of the wing colour and pattern 
to Polynesia, other morphological evidence suggests 
that Pseudopolynesia should be excluded from the 

Distribution. From Sundaland to New Guinea. 

Remarks. For further details see Holloway, 1997. 

Species included. One species. 

Pseudopolynesia amplificata Walker, 1861 


Pseudopolynesia hebe Bethune-Baker, 1915 

Pseudopolynesia Debauche 
Pseudoploynesia praelustris Prout 

New Guinea, Sulawesi, S. Moluccas. 

REMARKS. Holloway ( 1 997: 1 90) suggested that hebe 
and praelustris, which were previously described as 
subspecies of amplificata, might better be placed as 
'races' of hebe. 

Chaetolopha Warren, 1899 

(Fig. 186.) 

Chaetolopha Warren, 1 899: 4 1 . Type species: Scordylia 
oxyntis Meyrick, 1 89 1 : 8 1 7. 

The genus was included in the Asthenini by McQuillan 
& Edwards (in Nielsen, Edwards & Rangsi, 1 996: 227) 
and has been revised by Schmidt {in press). It includes 
an assemblage of species with considerable variation 
in genital structure. In none of these species does the 
morphology suggest that the genus belongs to 

Characters of the type species suggesting that 
Chaetolopha should excluded from Asthenini include 
a non-protuberant frons and well-developed, rough- 
scaled labial palpi. The uncus is well-developed. 
Structures resembling labides are well-sclerotized and 
arise from a pair of flask-shaped sclerites composing 
the juxta. We have not observed this rather peculiar 
arrangement in Asthenini. The sacculus of the valva is 
not extended. In the female genitalia, the signum is not 

Species included. Six species, all from Australia, 
are included by Schmidt (in press), who has assigned 
other species previously in Chaetolopha to a new 
genus from Papua New Guinea. 

Eleven species were listed in Scoble (1999). We 
have not reviewed the species, so they are not included 

Trichodezia Warren, 1895 

(Fig. 187.) 

Trichodezia Warren, 1895: 1 19. Type species: Odezia 
albovittata Guenee, [1858]: 520. 

Trichodezia was assigned to the Asthenini by Forbes 
(1948: 131) and listed there by Ferguson in Hodges 
(1983). Characters suggesting that this genus should 
be excluded from Asthenini include the presence of 


strong palpi and a well developed uncus. The ductus 
bursae is extremely short and the corpus bursae has a 
small signum, but not of the asthenine kind. 

A male and female of the type species (7^ albovittata) 
were examined. 

Distribution. North America, Japan, Eurasia. 

Species included. Five species were listed in Scoble 
(1999). We have not reviewed the species, so they are 
not listed here. 


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accentuata 95 

Acolutha 89 

adesma 88 

aedoea 102 

aeriferata 105 

aggerata 86 

Agnibesa 89 

ainoica 84 

albeolata 83 

albidaria 83 

albifera 84 

albinea 95 

albogilvaria 84 

albolucens 99. 100 

albosignata 83 

albovittata 109 

albulata 82, 83 

amblycoma 93 

amplificata 108, 109 

amurensis 83 

amylaria 106 

angularia 91 

anisobapta 93 

anseraria 83 

ansorgei 93 

anthodes 104 

Anydrelia 105 

aphrodite 96 

apicata 107 

apicistrigaria 95 

argentipuncta 84 

argyridia 94 

argyrorrhytes 84 

arizana 86 

Astenotricha 93 

Asthena 82 

Astheniodes 101 

Asthenotricha 93 

aurantiaca Prout ("Asthena") 85 

aurantiaca Hampson ( Hydrelia) 86 

Autallacta 85 

azela 99 

balausta 95 
baltoloma 102 
barnsae 93 
bella 86 
biangulata 95 
bicauliata 86 
bicolorata 86 
Bihastina 99 
binotata 86 
blomeri 96 
brevibasis 89 
brevipectinata 96 
brunneifasciata 84 
burmensis 102 

caeruleolineata 98, 99 
cambrica 95, 96 
cambricaria 96 
cambricata 96 
candace 94 
candidata 82, 83 

candidissima 84 
castaria 86 
centrata 87 
Chaetolopha 109 
Chalyboclydon 106 
'Chalyboclydon" 91 
character 102 
chibiana 87 
chionata 84 
cichisa 90 
cingulata 86 
cisseres 102 
Cleptocosmia 107 
comosissima 93 
comptaria 96 
condensata 86 
conisaria 96 
conspicuaria 86 
controversa 87 
corculina 83 
costalis 94 
crassisigna 96 
Cretheis 108 
crocearia 86 
curtitibia 105 
cymatodes 108 
cyparissaria 106 

deficiens 93 
dentatissima 93 
dharmsalae 105 
dilecta 96 
dimorpha 102 
Discoloxia 95 
disparata 94 
distinctaria 85 
distorta 105 
distrigaria 97 
duodecimlineata 97, 

elegans 87 
enisaria 87 
Eois 108 
Epicyme 90 
erutaria 96 
Eschatarchla 91 
euchloe 96 
Euchoeca 89 
euchroma 94 
eucosma 96 
euphorbiata 105, 10( 
euphylla 102 
eurychora 85 
euthecta 104 
exhumata 87 
expressa 97 

femandi 93 
flammeolaria 87 
flammulata 87 
flavicata 87 
flavicoma 93 
flavidula 88 
flavilinea 87 

flavostrigata 87 

flexilinea ('Chalyboclydon') 91, 92 

flexilinea (Parasthena) 101 

formosana 9 1 

fragilis 103 

fulgurata 88 

furtiva 93 

fuscata 106 

fuscocastanea 87 

gemmifera 99 
glaucosa 103 
gracilipennis 87 
grandis 93 

hamadryas 83 
Hastina 98 
hebe 109 
hepararia 89, 90 
Hippolyte 90 
Horisme 80 
Hydrelia 85 
hypoconia 96 

ilara 96 
impleta 87 
inefficax 96 
inhaesa 103 
inornata 87 
inutilis 94 
iopolia 103 
ischnophrica 103 
italicata 106 

japonica 87 

kasyata 96 
kioudjrouaria 96 
kukunoora 97 

lactularia 83 
laetivirga 87 
laria 96 
lassa 83 
latsaria 87 

Leucoctenorrhoe 100 
leucogramma 87 
leucydra 103 
lilacina 96 
limata 96 
limburgia 106 
limnaea 103 

lineata (Eschatarchia) 91 
lineata (Hydrelia) 87 
lineata (Venusia) 96 
livida 83 
lophopterata 94 
lucata 87 
lutea 106 
luteata 87 
luteosparsata 87 

Macrohastina 99 
malostigma 94 
maniata 96 



marginata 107 
marginepunctata 87 
marmoraria 96 
megaspilata 97 
melanogramma 96 
melanosticta 84 
mera 100 
meroplyta 89 
meruana 94 
microgyria 103 
microptera 87 
Minoa 105 
mionoseista 95 
mullata 91 
murinata 106 
musculata 87 
mutabilis 107, 108 

naparia 97 
nebulata 89, 90 
nebulosaria 96 
nepalensis 87 
nesiotes 94 
nigrifurca 97 
nisaria 87 
Nomenia 97 
nubivaga 103 
nymphaeata 84 

obliquisigna 95, 97 
obliterata 90 
obsoleta 98 
oceanias 103 
ochrearia 87 
ochrifasciaria 84 
ochrota 97 
octomacularia 84 
opedogramma 84 
ornata 87 
oxyntis 109 

pallidaria 97 
Palpoctenidia 92 
palumbes 96 
pampesia 88 
panapala 103 
papuensis 104 
parabolica 94 
paradoxa 97 
Parasthena 101 
participata 97 
parvularia 87 
parvulata 87 
paucilinea 103 
pavonica 87 
pearsalli 97 
pellucida 103 
percandidata 84 
perlineata 96 
phaeodryas 103 
phanoides 109 

phasma 97 

phoenicosoma 92 

pictaria 89 

pisicolor 103 

planicaput 97 

plenaria 84 

pleopictaria 89 

plicataria 105 

plumbeolineata 89 

pluristrigata 99 

Poecilasthena 101 

polycymaria 101, 103 

polydora 94 

Polynesia 104 

praelustris 109 

proschora 94 

prouti 103 

psephotaenia 94 

Pseudopolynesia 108 

Pseudostegania 80 

pulchraria (Poecilasthena) 101. 103 

pulchraria (Venusia) 96 

punctilinearia 89 

punctiuncula 97 

purpuraria 97 

pycnoconia 94 

pythia 94 

quadrata 94 
quadrilinea 100 
quadripunctata 84 

rala 96 

recurvilineata 89 
rhodoptera 88 
risata 91 
Roessleria 83 
roseicosta 96 
rubraria 88 
rubricosta 88 
rubrilinea 88 
rubrivena 88 
rubropunctaria 90, 91 
rufigrisea 88 
rufinota 88 
russearia 108 

sachaliensis 84 
sachalinensis 88 
salienta 96 
sanguiflua 88 
sanguiniplaga 88 
schistaria 104 
scitula 97 
scitularia 96 
scoliota 103 
scotozona 88 
secunda 98 
semidivisa 94 
semilauta 92 
semistrigata 97 

sencea 88 
serraticomis 94 
shioyana 88 
shuotsu 96 
sikkimensis 97 
sinuosata 87 
sjostedti 94 
soldaria 83 
sordiata 106 
speciosa 88 
stenozona 99 
Sterrhochaeta 80 
sthenommata 103 
straba 94 
straminearia 85 
strangulata 95 
strigata 90 
subcingulata 88 
subditaria 85 
subfalcaria 99 
sublatsaria 88 
subobliquaria 88 
subpurpureata 103 
subtestacea 88 
subviridata 100 
sunandava 104, 105 
sylvata 88 
syngenes 97 
szechuanensis 96 

tamsi 95 
tchraria 97 
tchratchraria 84 
tenera 88 
terraenovae 88 
testaceata 88 
thalassias 103 
torata 94 
Trichodezia 109 
triseriata 84 
tripogonias 94 
truncapex 104. 105 
tuhuata 103 

ulula 88 
undularia 88 
undulata 84 
undulosata 88 
unicolorata 106 
unipecten 95 
urarcha 103 

Venusia 95 
venusta 89 
vexata 91 
violettaria 97 
viridata 100 

xylocyma 104 

yargongaria 85 
yasudai 97 

1 A. albidaria 

5 A. lassa 


2 A. albosignata 

6 A. Uvida 

3 A. albulata 

■ -t 

7 A. melanosticta 


^ I >J 

4 A. anseraria anseraria 

8 A. nymphaeata 

1 ' V'""' 

9 A. ochrifasciaria 


13 A. sachaliensis 

17 H. arizana 

I. bicolorata 

10 A. octomacularia 

14 A. tchratchraria 

18 H. aurantiaca 

11 A. opedogramma 12 A. p/enaria 

&M0 f * f 

. 1 ■...-; 

4^fS : - 

■ g ' - 

22 H. binotata 

15 A. undulata 

16 H. aggerata 

19 H. be//a 

20 H. bicauliata 

23 H. castaria 

24 H. cingulata 

Figs 1-24. Adults. 1-15, Asthena spp.; 16-24, Hydrelia spp. Scale lines: 10 mm. 

25 ft condensate 

26 ft. conspicuaria 

27 ft. controversa 

28 ft crocearia 

29 ft elegans 

30 ft enisaria 

3 1 ft flammeolaria 

32 ft flavilinea 

33 ft fuscocastenea 

34 ft gracilipennis 

37 ft laetivirga 

38 ft leucogramma 

41 ft marginepuncteta 42 ft microptera 

,$ ~- : 

39 ft iineata 

40 ft /ucata 

43 ft nepalensis 

44 ft nisaria 

45 ft ochrearia 

46 ft ornate 

47 ft parvularia 

48 ft parvulate 

Figs 25-48. Adults. Hydrelia spp. Scale lines: 10 mm. 

49 H. rhodoptera 

53 H. rubrivena 

61 H. speciosa 

50 H. rubraria 

5 1 H. rubricosta 

54 H. mfigrisea 

55 H. rufinota 

56 H. sanguiflua 

57 H. sanguiniplaga 


58 H. sericea sericea 59 H. sericea pampesia 60 H. shioyana 

" Vsy 

62 H. subcingulata 63 H. sublatsaria 

64 H. subobliquaria 

65 H. subfestacea 

66 H. sy/ va fa 

tf %w# 

67 H. undularia 

68 H. undulosata 

'•> *w" 

- - • *v > - 




' | 

69 A. pictaria pictaria 70 A. pictaria brevibasis 71 A. plumbeolineata 72 A. punctilinearia 

Figs 49-72. Adults. 49-68, Hydrelia spp.; 69-72, Agnibesa spp. Scale lines: 10 mm. 


73 A. recurvilineata 

74 A. recurvilineata 

75 A. venusta 

76 E. nebulata 

77 E. rubropunctaria 78 E. lineata 

79 "C. " flexilinea 

80 P. phoenicosoma 

81 P. phoenicosoma 

82 A. amblycoma 

83 A. anisobapta 

84 A. ansoTgei 

85 A. bamsae 

86 A. dentatissima 

87 A. femandi 

88 A. flavicoma 

89 A. /urti'va 

90 A. lophopterata 

91 A. malostigma 

92 A. nesiofes 

93 A. polydora 

94 A. proschora 

95 psephotaenia 

96 A. pycnoconia 

Figs 73-96. Adults. 73-75, Agnibesa spp.; 76, Euchoeca spp.; 77, Epicyme sp.; 78, Eschatarchia sp.; 79, ' Chalyboclydon ' 
sp.; 80-81, Palpoctenidia phoenicosoma; 82-96, Asthenotricha spp. Scale lines: 10 mm. 

I N. 



97 A. pythia 

98 A. semidivisa semidivisa 99 A. semidivisa euchroma 100 A. serraticomis 


W/1 w 

101 A. straba 

102 A. torata 103 A. tripogonias 

104 A. argyridia 

106 A. costalis 

107 A. inutj/is 

108 A. meruana 

109 A. sjostedti sjostedti 

^ 4 * 


113 V.albinea 

114 V. blomeri blomeri 115 V brevipectinata 116 V cambrica cambrica 

117 V comptaria 

118 V conisaria conisaria 119 V crassisigna 

120 V eucosma 

Figs 97-120. Adults. 97-1 11, Asthenotricha spp.; 1 12-120, Vfe/n«/a spp. Scale lines: 10 mm. 

.:;% JK 


•- '■■Wm$-'''t 



V V ■ 


V inefficax 

7 '¥ ■ . i 

^M 1 V_. 

122 V. kioudjrouaria 123 V iaria laria 

124 V lilacina lilacina 


125 V.limata 

126 V iineafa 

127 V marmoraria 128 V megaspilata 

*>W ^^r^HK^ ' * X/'"-" 

,. 1<fi 


129 Vm'grifiirca 

130 V. obliquisigna 


131 Vochrota 

132 V. pallidaria 



. y 

133 Vpiiasma 


137 V. semistrigata 

141 N. duodecimlineata 

134 V.planicaput 

135 V punctiuncula 136 VCpiupuraria 

138 V sikkimensis 

139 V fdiraria 

142 H. caemleolineata 143 H. pluristrigata 

140 Vyasudai 




*vS 1 ' ^1" 


144 M. aze/a 

Figs 121-144. Adults. 121-140, Venusia spp.; 141, Nomenia sp.; 142-143, Hastina spp.; 144, Macrohastina sp. Scale lines: 
10 mm. 

145 M. gemmifera 

146 M. stenozona 

149 B. viridata 

150 L. quadrilinea 

! F^s. 

' 1 

153 P balioloma 

157 P euphylla 


154 P burmensis 

158 P inhaesa 

.■ A 

147 B. albolucens 

148 B. subviridata 

151 P flexilinea 

152 P aedoea 

' \/ 


155 P. character 

156 P. dimorpba 


159 P. leucydra 160 P. limnaea 


161 P. nubivaga 

162 P paucilinea 

163 P.prouti 

165 P scoliota 

1 64 P. pulchraria 



166 P subpuipureata 167 P eufhecta 

168 P. papuensis 

Figs 145-168. Adults. 145-146. Macrohastina spp.; 147-149, Bihastina spp.; 150, Leucotenorrhoe sp.; 151, Parasthena 
sp.; 152-168, Poecilasthena spp. Scale lines: 10 mm. 



169 P. anthodes 

170 P. schistaria 

171 P.xylocyma 


* 1 * ?> . 

172 P. curtitibia 

173 P. sunandava 

177 A. plicataria 

181 C.wutabilis 

174 P. tnmcapex 

175 A. dharmsalae 

185 Pfiebe 

178 M. murinata murinata 179 Minoasp. 

182 E. cymatodes 

183 E.plicata 

186 C. oxynfis 

187 T. albovittata 

176 A. distorta 

1 84 E. russearia 

188 S. Mgurata 

Figs 169-188. 169-171, Poecilasthena spp.; 172-174, Polynesia spp.; 175-177, Anwfre/ra spp.: 178-179, M;'«oa spp.; 180, 
Chalyboclydon sp.; 181, Cleptocosmia sp.; 182-184, £ow spp.; 185, Pseudopolynesia sp.; 186, Chaetolopha sp.; 187, 
Trichodezia sp., 188, Sterrhochaeta sp. Scale lines: 10 mm. 



Figs 189-192. Wing venation. 189, Asthena albulata; 190. Agnibesa pictaria; 191. Venusia cambrica; 192, Hastina 
caerulineata. Scale lines: 189, 192, 1.0 mm.; 190, 191, 2.5 mm. 

193 A. albulata 

194 A. opedogramma 

195 A. undulaia 

196 H. beila 


, <^ 

197 H.sylvata 

198 A. pictaria 

Figs 193-198. Male genitalia. 193-195, Asthena spp.; 196-197, Hydrelia spp.; 198, Agnibesa sp. 

199 E. nebulata 

200 E. rubropunctaria 

v y 

201 E. Imcaiu 

202 "C. " flexilinea 

203 P. pboenicosoma semilauta 

204 A. dentatissima 

Figs 199-204. Male genitalia. 199, Euchoeca sp.; 200, Epicyme sp.; 201. Eschatarchia sp.; 202, 'Chalyboclydon' sp.; 203, 
Palpoctenidia sp.; 204. Asthenotricha sp. 


205 A. argyridia 

206 V cambrics 

209 N. duodecimlineata 

210 H. cacrukolincata 

Figs 205-210. Male genitalia. 205, Asthenotricha sp.; 206-208, Venusia spp.; 209, Nomenia sp.; 210, Hastina sp. 

211 M. azela 

212 B. alboluccns 

, < V, 


213 L. quadrilinea 

214 P. flexilinea 

215 P. dimorpha 


Figs 211-215. Male genitalia. 21 1, Macrohastina sp.; 212. Bihastina sp., 213, Leucoctenorrhoe sp.; 214. Parasthena sp.: 
215, Poecilasthena sp. 

216 P. pulchraria 


217 P. papuensis 

218 P. schislaria 

219 P. sunandava 


220 A. plicataria 

221 M. murinata 

Figs 216-221. Male genitalia. 216-218, Poecilasthena spp., 219, Polynesia sp.; 220, Anydrelia sp.; 221, Minoa sp. 




222 C. marginals 

223 C mutabilis 

224 E. cymatodes 

225 P.hebc 

Figs 222-225. Male genitalia. 222, Chalyboclydon sp.; 223, Cleptocosmia sp.; 224, Eois sp.; 225, Pseudopolynesia sp. 

226 A. albulata 227 A. opedogramma 228 A. undulaia ll 1 ) H. bella 

230 H. sylvata 



231 A. pictaria 232 E. nebulata 233 E. rubropunctaria 234 E. lineata 235 "C." flexilinca 

236 P. phocnicosoma 237 A. dcntatissima 238 A. argyridia 

239 V cambrica 240 V obliquisigna 

Figs 226-240. Aedeagus. 225-228, Asthenct spp.; 229-230, Hydrelia spp.; 231, Agnibesa sp.; 232, Euchoeca sp.; 233, 
Epicyme sp.; 234, Eschatarchia sp.; 235, 'Chalyboclydon sp.; 236, Palpoctenidia sp.; 237-238. Asthenotricha spp.; 239- 
240, Venusia sp. 



241 V. ochrota 242 N. duodccimlineata 243 H. caerulcolincata 244 M. aze/a 245 B. alboluccns 




246 L. quadrilinca 247 P. flexilinea 248 P. dimorpha 249 P. pukhraria 250 P papuensis 

251 P. sehisfaria 252 P. sunandava 253 A. plicataria 

Figs 241-253. Aedeagus. 241, Ve/it/s/a sp.; 242, Nomenia sp.; 243, Hastina sp.; 244, Macrohastina sp.; 245, Bihastina sp. 
246, Leucoctenorrhoe sp.; 247, Parasthena sp; 248-25 1 , Poecilasthena spp.; 252, Polynesia sp.; 253, Anydrelia sp. 

254 M. murirtata 255 C. marginals 256 C. mutabilis 257 E. cymatodes 258 P. hcbe 

• ' 

• •'""'■fop 
,,„, — « ■p*i-\ 


259 P. phocnicosoma 260 P. flexilinca 

261 P. Hexilinea 

262 P. dimorpha 

263 P. pukhraria 

264 P. papuensis 

265 P. schistaria 

266 P. suna/idava 

267 M. rnunnala 

Figs 254-267. 254-258, aedeagus; 259-267, male abdomen. 254, Minoa sp.; 255, Chalyboclydon sp.; 256, Cleptocosmia 
sp.; 257, £b/.s' sp.; 258, Pseudopolynesia sp.; 259, Palpoctenidia sp.; 260-261, 260 sternum, 261 tergum, Parasthena sp.; 
262-265, Poecilasthena spp.; 266, Polynesia sp.; 267. Minoa sp. 


268 A.albulata 

272 £ ncbulata 

276 P. phoenicosoma 

w ,y 

269 A. tchratchraria 


%^-r f 

270 H. bella 

273 E. rubronuncfaria 274 E. lincata 

271 A. pictaria 

275 "C." flcxilinca 

277 A. dcntatissima 

278 A. argyridia 

279 V cambrica 

Figs 268-279. Female genitalia. 268-269, Asthena spp.; 270. Hydrelia sp.; 21 \ , Agnibesa sp.; 272, Euchoeca sp.; 273, 
Epicyme sp.; 274, Eschatarchia sp.; 275, 'Chalyboclydon' sp.; 276. Palpoctenidia sp.; 277-278, Asthenotricha sp.; 279, 
Venusia sp. 

280 V. obliquisigna 

281 N. duodecimlineata 282 H. cacruleolineata 

283 M. azcla 



284 B. alboluccns 

285 L. quadrilinca 

286 P. llcxilinea 

287 P. dimorpha 

288 P. pukhraria 

289 P. papuensis 

290 P. schistaria 

291 P. sunandava 

Figs 280-291. Female genitalia. 280, Venusia sp.: 2&\,Nomenia sp.; 282, Hastina sp.; 283, Macrohastina sp.; 284. 
Bihastina sp.; 285, Leucoctenorrhoe sp.; 286, Parasthena sp.; 287-290, Poecilasthena spp.; 291, Polynesia sp. 





292 A. plicataria 293 M. munnaia 

294 C marginata 

295 P. dimoq)ha 

Figs 292-295. Female genitalia. 292, Anydrelia sp.; 293, Minoa sp.: 294, Chalyboclydon sp.; 295, signum of 
Poecilasthena sp. 

Bulletin of The Natural History Museum 
Entomology Series 

Earlier Entomology Bulletins are still in print. The following can be ordered from Intercept (address on inside front 
cover). Where the complete backlist is not shown, this may also be obtained from the same address. 

Volume 49 

No. 1 Afrotropical jumping plant lice of the family Triozidae (Homoptera: Psylloidea-. D. Hollis. 1984. Pp. 1- 

102, 324 tigs. £15.30 

No. 2 The taxonomy of the western European grasshoppers of the genus Euchorthippus, with special reference 

to their songs (Orthoptera: Acrididae). D.R. Ragge & W.J. Reynolds. 1984. Pp. 103-151. 88 figs. £7.20 
No. 3 An historical review of the higher classification of the Noctuidae (Lepidoptera). I.J. Kitching. 1984. Pp. 

153-234, 4 figs. £12.00 

No. 4 The Pimplinae, Xoridinae, Acaenitinae and Lycorininae (Hymenoptera: Ichneumonidae) of Australia. I.D. 

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No. 5 The Palaearctic species of Ascogaster (Hymenoptera: Braconidae). T Huddleston. 1984. Pp. 341-392. 79 

figs. £7.80 

Volume 50 

No. 1 Taxonomy of Neotropical Derbidae in the new tribe Mysidiini (Homoptera). PS. Broomfield. 1985. Pp. 1- 

152.501 figs. £22.80 

No. 2 Nymphal taxonomy and systematica of the Psylloidea (Homoptera). l.M. White & I.D. Hodkinson. 1985. 
Pp. 153-301, 201 figs, 18 tables. £23.00 

No. 3 The Whitefly of New Guinea (Homoptera: Aleyrodidae). J.H. Martin. 1 985. Pp. 303-35 1 , 48 figs. £8.00 

Volume 51 

No. I The ichneumon-fly genus Banchus (Hymenoptera) in the Old World. M.G. Fitton. 1985. Pp. 1-60, 129 

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No. 2 The phylogeny, classification and evolution of parasitic wasps of the subfamily Ophioninae 

(Ichneumonidae). I.D. Gauld. 1985. Pp. 61-185, 52 figs. £21.00 

No. 3 A cladistic analysis and classification of trichodeclid mammal lice (Phthiraptera: Ischnocera). C.H.C. Lyal. 

1985. Pp. 187-346. 250 figs. £26.00 

No. 4 The British and some other European Eriococcidae (Homoptera: Coccoidea). D.J. Williams. 1985. Pp. 

347-393. 1 8 figs. £8.00 

Volume 52 

No. 1 The sandflies of Egypt (Diptera: Phlebotominae). R.P Lane. 1986. Pp. 1-35. 80 figs. 4 tables. £5.60 

No. 2 Fungus moths: a review of the Scardiinae (Lepidoptera: Tineidae). G.S. Robinson. 1986. Pp. 37-181. 200 

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No. 3 A revision of the European Agathidinae (Hymenoptera: Braconidae). G.E.J. Nixon. 1986. Pp. 183-242, 68 

figs. £11.00 

No. 4 A key to the Afrotropical genera of Eucoilidae (Hymenoptera) with a revision of certain genera. J. 

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Volume 53 

No. 1 A review of Miletini (Lepidoptera: Lycaenidae). J.N. Eliot. 1986. Pp. 1-105, 108 figs. £18.00 

No. 2 Australian ichneumonids of the tribes Labenini and Poecilocryptini. I.D. Gauld & G.A. Holloway. 1986. 

Pp. 107-149, 65 figs. £8.40 

No. 3 The tribe Pseudophloeini (Hemiptera: Coreidae) in the Old World tropics with a discussion on the 

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No. 4 The songs of the western European grasshoppers of the genus Omocestus in relation to their taxonomy 

(Orthoptera: Acrididae). D.R. Ragge. 1986. Pp. 213-249, 128 figs. £7.50 

No. 5 The structure and affinities of the Hedyloidea: a new concept of the butterflies. M.J. Scoble. 1986. Pp. 

251-286, 102 figs. £7.00 

Volume 54 

No. 1 Studies on the Old World species of Holothrips (Thysanoptera: Phlaeothripidae). S. Okajima. 1987. Pp. 1- 

74, 207 figs. £14.00 

No. 2 Spectacles and Silver Ys: a synthesis of the systematics, cladistics and biology of the Plusiinae (Lepidop- 
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No. 3 A review of the Solenopsis genus-group and revision of Afrotropical Monomorium Mayr (Hymenoptera: 

Formicidae). B. Bolton. 1987. Pp. 263-452, 100 figs. £36.50 

Volume 55 

No. 1 A reclassification of the European Tetrastichinae (Hymenoptera: Eulophidae), with a revision of certain 

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No. 2 The songs of the western European grasshoppers of the genus Stenoboihrus in relation to their taxonomy 

(Orthoptera: Acrididae). D.R. Ragge. 1987. Pp. 393-424, 1 16 figs. £6.00 

Volume 56 

No. 1 The legume-feeding psyllids (Homoptera) of the wet Palaearctic Region. I.D. Hodkinson & D. Hollis. 

1987. Pp. 1-86, 294 figs. £16.00 

No. 2 A review of the Malvales-feeding psyllid family Carsidaridae (Homoptera). D. Hollis. 1987. Pp. 87-127, 

94 figs. £6.00 

No. 3 A review of the Rhadalinae (=Aplocneminae) (Coleoptera: Melyridae). E.R. Peacock. 1987. Pp. 129-170, 

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No. 4 A revision of some Afrotropical genera of Eucoilidae (Hymenoptera). J. Quinlan. 1988. Pp. 171-229. 199 

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Volume 57 

No. 1 A survey of the Ophioninae (Hymenoptera: Ichneumonidae) of tropical Mesoamerica with special 

reference to the fauna of Costa Rica. I.D. Gauld. 1988. Pp. 1-309, 352 figs, 32 maps. £52.00 

No. 2 A taxonomic revision of Alabagrus (Hymenoptera: Braconidae). M.J. Sharkey. 1988. Pp. 31 1-437, 28 

figs, 22 maps. £24.50 

No. 3 A taxonomic revision of Caryocolum (Lepidoptera: Gelechiidae). P. Huemer. 1988. Pp. 439-571, 221 figs. 


Volume 58 

No. 1 The mealybug genus Plcmococcus (Homoptera: Pseudococcidae). J.M. Cox. 1989. Pp. 1-78, 40 figs. 

No. 2 The Simuliidae (Diptera) of the Santiago onchocerciasis focus of Ecuador. A.J. Shelley, M. Arzube & C.A. 
Couch. 1989. Pp. 79-130. 153 figs (including 2 plates in colour). 

Volume 59 

No. 1 The songs of the western European bush-crickets of the genus Platycleis in relation to their taxonomy 

(Orthoptera: Tettigoniidae). D.R. Ragge. 1990. Pp. 1-35. 

A reclassification of the Melanotus group of genera (Coleoptera: Elateridae). C.M.F. von Hayek. 1990. 

Pp. 37-115. 
No. 2 The green lacewings of the world: a generic review (Neuroptera: Chrysopidae). S.J. Brooks & PC. 

Barnard. 1990. Pp. 117-286. 

Volume 60 

No. 1 The bumble bees of the Kashmir Himalaya (Hymenoptera: Apidae, Bombini). P.H. Williams. 1991. 

Pp. 1-204. 
No. 2 Sattleria: a European genus of brachypterous alpine moths (Lepidoptera: Gelechiidae). L.M. Pitkin & K. 

Sattler. 1991. Pp. 205-241. 

A review of wing reduction in Lepidoptera. K. Sattler. 1991. Pp. 243-288. 

Volume 61 

No. 1 Thrips (Thysanoptera) from Pakistan to the Pacific: a review. J.M. Palmer. 1992. Pp. 1-76. 

No. 2 Neotropical red-brown Ennominae in the genera Thysanopyga Herrich-Scaffer and Perissopteryx Warren 
(Lepidoptera: Geometridae). M. Kruger & M.J. Scoble. 1992. Pp. 77-148. 

Volume 62 

No. 1 Caloptilia leaf-miner moths (Gracillariidae) of South-East Asia. Decheng Yuan and Gaden S. Robinson. 

1993. Pp. 1-37. 
No. 2 Neotropical Emerald moths of the genera Nemoria, Lissochlora and Chavarriella, with particular 

reference to the species of Costa Rica (Lepidoptera: Geometridae. Geometrinae). Linda M. Pitkin. 1993. 

Pp. 39-159. 

Volume 63 

No. 1 A revision of the Indo-Pacific species of Ooencyrtus (Hymenoptera: Encyrtidae), parasitoids of the 

immature stages of economically important insect species (mainly Hemiptera and Lepidoptera). D.W. 
Huang and J.S. Noyes. Pp. 1-135. 

No. 2 A taxonomic review of the common green lacewing genus Chrysoperla (Neuroptera: Chrysopidae). S.J. 
Brooks. Pp. 137-210. 

Volume 64 

No. 1 Revision of the neotropical genus Oospila Warren (Lepidoptera: Geometridae) M. A. Cook and M.J. 

Scoble. Pp. 1-115. 
No. 2 Encyrtidae of Costa Rica (Hymenhoptera: Chalcidoidea): the genus Aenasius Walker, parasitoids of 

mealybugs (Homoptera: Pseudococcidae). J.S. Noyes and H. Ren. Pp. 1 17-164. 

Volume 65 

No. 1 A revised classification of the Asian and Pacific selenocephaline leafhoppers (Homoptera: Cicadellidae). 

Y. Zhang and M.D. Webb. Pp 1-103. 
No. 2 Encyrtidae (Hymenoptera: Chalcidoidea) of Costa Rica: the genera and species associated with jumping 

plant-lice (Homoptera: Psylloidea). J.S. Noyes and P. Hanson. Pp. 105-164. 

Volume 66 

No. 1 Biosystematic studies on the Simuliidae (Diptera) of the Amazonia onchocerciasis focus. A.J. Shelley, 

C.A. Lowry, M. Maia-Herzog, A. P. A. Luna Dias and M.A.P Moraes. Pp. 121. 
No. 2 Microtermes in East Africa (Isoptera: Termitidae: Macrotermitinae) S. Bacchus. 1997. Pp. 123-171. 

Volume 67 

No. 1 Mealybugs of the genera Eumyrmococcus Silvestri and Xenococcus Silvestri associated with the ant genus 

Acropyga Roger and a review of the subfamily Rhizoecinae (Hemiptera, Coccoidea, Pseudococcidae). D.J. 

Williams. 1998. Pp. 1-64. 

Monophyly of the dacetonine tribe-group and its component tribes (Hymenoptera: Formicidae). B. Bolton. 

1998. Pp. 65-78. 

An annotated checklist of bumble bees with an analysis of patterns of description (Hymenoptera: Apidae, 

Bombini). PH. Williams. 1998. Pp. 79-152. 
No. 2 A revision of the leafhopper tribe Paraboloponini (Hemiptera: Cicadellidae: Selenocephalinae) in the 

Indian subcontinent. C.A. Viraktamath. 1998. Pp. 153-208. 

A review of the Palaearctic Neorhacodinae (Hymenoptera, Ichneumonidae) with Eremura Kasparyan, 

1995 new to the west Palaearctic. D.G. Notton and M.R. Shaw. 1998. Pp. 209-218. 

Volume 68 

No. 1 Revision of the Oriental Opostegidae (Lepidoptera) with general comments on phytogeny within the 

family. Rimantas Puplesis and Gaden S. Robinson. 1999. Pp. 1-92. 
No. 2 A revision of the African and Malagasy species of the genus Leptomastix (Hymenoptera, Encyrtidae), 

parasitoids of mealybugs (Homoptera: Pseudococcidae). Jean-Marc Anga and John S. Noyes. 1999. Pp. 


A revision of the north-west European species of the formosus species group of Spihmicrus (Hymenop- 
tera: Diapriidae). David G. Notton. 1999. Pp. 129-144. 

A review of the soldierless African termite genus Amicotermes Sands 1972 (Isoptera, Termitidae, 

Apicotermitinae). W.A. Sands. 1999. Pp, 145-193. 

Volume 69 

No. 1 Editorial. R.I. Vane- Wright. 2000. Pp. 1-2. 

A review of the Central and South American Nepticulidae (Lepidoptera) with special reference to Belize. 

Rimantas Puplesis and Gaden S. Robinson. 2000. Pp. 3-1 13. 
No. 2 Systematics of Onirion, a new genus of Sabethini (Diptera: Culicidae) from the Neotropical Region. R.E. 

Harbach and E.L. Peyton. 2000. Pp. 115-169. 

The Simuliidae (Diptera) of the secondary onchocerciasis focus at Minacu in central Brazil. A.J. Shelley. 

M. Maia-Herzog, C.A. Lowry, A.P.A. Luna Dias, PR. Garritano, A. Shelley, M. Camargo, H.G. Carter. 

2000. Pp. 171-221. 

Volume 70 

No. 1 A revision of the tribe Macariini (Lepidoptera: Geometridae: Ennominae) of Africa, Madagascar and 

Arabia. Martin Kriiger. 2001. Pp. 1-502. 
No. 2 A revision of the afrotropical species of the genus Graphium (Lepidoptera: Rhopalocera: Papilionidae). 

Campbell R. Smith and R.I. Vane-Wright. 2001. Pp. 503-719. 


1 Loboponera gen.n. and a review of the Afrotropical Plectroctena genus 
group (Hymenoptera: Formicidae) 

Barry Bolton and William L Brown, Jr. 

19 New Neotropical Nepticulidae (Lepidoptera) from the western 
Amazonian rainforest and the Andes of Ecuador 
Rimantas Puplesis, Arunas DiSkus and Gaden S. Robinson 

59 A review and checklist of the Neotropical Nepticulidae (Lepidoptera) 
Rimantas Puplesis, Arunas DiSkus, Gaden S. Robinson and Giovanni Onore 

79 A review of the genera associated with the tribe Asthenini (Lepidop- 
tera: Geometridae: Larentiinae) 
Dayong Xue and Malcolm J. Scoble 


Vol.71, No. 1, June 2002